Circadian rhythms enable efficient resource selection in a human‐modified landscape

Animals access resources such as food and shelter, and acquiring these resources has varying risks and benefits, depending on the suitability of the landscape. Some animals change their patterns of resource selection in space and time to optimize the trade‐off between risks and benefits. We examine the circadian variation in resource selection of swamp wallabies (Wallabia bicolor) within a human‐modified landscape, an environment of varying suitability. We used GPS data from 48 swamp wallabies to compare the use of landscape features such as woodland and scrub, housing estates, farmland, coastal areas, wetlands, waterbodies, and roads to their availability using generalized linear mixed models. We investigated which features were selected by wallabies and determined whether the distance to different landscape features changed, depending on the time of the day. During the day, wallabies were more likely to be found within or near natural landscape features such as woodlands and scrub, wetlands, and coastal vegetation, while avoiding landscape features that may be perceived as more risky (roads, housing, waterbodies, and farmland), but those features were selected more at night. Finally, we mapped our results to predict habitat suitability for swamp wallabies in human‐modified landscapes. We showed that wallabies living in a human‐modified landscape selected different landscape features during day or night. Changing circadian patterns of resource selection might enhance the persistence of species in landscapes where resources are fragmented and disturbed.     

Some Consequences of Size Variability in Juvenile Prickly Sculpin, Cottus asper

Variation in size among fishes can have important ecological consequences. The origin and persistence of variation in size among a cohort of juvenile prickly sculpin, Cottus asper, was explored experimentally by manipulating the size variability in juvenile fish and the presence of conspecific adult fishes in a factorial design. I found that adult fish significantly suppressed the growth and survivorship of juveniles, while the variance treatment alone had no effect on growth or survivorship. The presence of adults was also associated with a decrease in size variability in the high variance treatment. An analysis of juvenile growth patterns revealed no evidence for size dependent growth or temporal correlations in growth (`growth autocorrelation'), and are consistent with a lack of amplification of size variance through time. Juvenile C. asper appear to have been feeding on a relatively homogeneous resource and utilizing those resources similarly. Thus, patterns of individual growth and the presence of adult conspecifics may both serve to dampen size differences in this coastal fish species.     

Competitive Outcomes of Aquatic Container Diptera Depend on Predation and Resource Levels

Resources and predation are both known to be important in structuring communities; however the strength of one factor may be affected by the intensity of the other. This study used a fully crossed factorial experiment in laboratory microcosms to examine the ability of a predator, Corethrella appendiculata (Grabham), and basal resources (leaf litter) to differentially affect two competing species of mosquito prey. Increased resources resulted in shorter developmental time and increased survivorship, mass, and population performance for both prey species, except when predation levels were high. Increased levels of predation and resources reduced the negative competitive effects of Aedes albopictus (Skuse) on Ochlerotatus triseriatus (Say). At low levels of resources and predation, the superior competitor, A. albopictus had the higher survivorship, and at high levels of resources and predation, the inferior competitor's survival was greater. Predators in high-resource treatments emerged larger than those in low resources, suggesting the occurrence of a bottom-up cascade or alternative feeding method. This study suggests that survival and coexistence of the two prey species may depend on the interaction of resources and predation, in that high levels of predation are important for the coexistence of both species.     

Influence of resource levels,organic compounds and laboratory colonization on interspecific competition between the Asian tiger mosquito Aedes albopictus (Stegomyia albopicta) and the southern house mosquito Culex quinquefasciatus

The mosquitoes Aedes albopictus (Stegomyia albopicta) (Skuse) and Culex quinquefasciatus (Say) (Diptera: Culicidae) are common inhabitants of tyres and other artificial containers, which constitute important peridomestic mosquito breeding habitats. We tested the hypotheses that interspecific resource competition between the larvae of these species is asymmetrical, that the concentration of chemicals associated with decomposing detritus affects the competitive outcomes of these species, and that wild and colonized strains of Cx. quinquefasciatus are affected differently by competition with Ae. albopictus. We conducted two laboratory competition experiments wherein we measured survivorship and estimated population growth (λ′) in both species under multiple mixed‐species densities. Under varying resource levels, competition was asymmetrical: Ae. albopictus caused competitive reductions or exclusions of Cx. quinquefasciatus under conditions of limited resources. In a second experiment, which used both wild and colonized strains of Cx. quinquefasciatus, organic chemical compounds associated with decomposing detritus did not affect the competitive outcome. The colonized strain of Cx. quinquefasciatus had greater survivorship and adult mass, and faster development times than the wild strain, but both strains were similarly affected by competition with Ae. albopictus. Competition between these species may have important consequences for vector population dynamics, especially in areas in which tyres and artificial containers constitute the majority of mosquito breeding habitats.     

EFFECTS OF LIGHT AND DENSITY ON RESOURCE ALLOCATION IN A FOREST HERB,ASTER ACUMINATUS (COMPOSITAE)

Recent studies show that forest herbs have low sexual reproductive efforts (SRE) and high vegetative reproductive efforts (VRE) and that a variety of factors may affect the levels of reproduction and the distribution patterns of forest species. Aster acuminatus is a micro-successional forest herb common in northern New England. Resource allocation patterns were determined for seven aster patches varying in density and light level. Plant density affects neither plant size nor any aspect of resource allocation, and VRE remains constant over all density and light levels. Light level is, however, significantly correlated with patch location, average plant size and average patch SRE. Only large plants produce flowers suggesting that light affects sexual reproduction indirectly through effects on plant size. The high rhizome and low seed survivorship of this species may partly explain the constant VRE and varying SRE levels, although the resource allocation patterns observed do not correspond well with the predictions of certain theoretical models.     

Response of a specialist bat to the loss of a critical resource

Human activities have negatively impacted many species, particularly those with unique traits that restrict their use of resources and conditions to specific habitats. Unfortunately, few studies have been able to isolate the individual and combined effects of different threats on population persistence in a natural setting, since not all organisms can be associated with discrete habitat features occurring over limited spatial scales. We present the results of a field study that examines the short-term effects of roost loss in a specialist bat using a conspicuous, easily modified resource. We mimicked roost loss in the natural habitat and monitored individuals before and after the perturbation to determine patterns of resource use, spatial movements, and group stability. Our study focused on the disc-winged bat Thyroptera tricolor, a species highly morphologically specialized for roosting in the developing furled leaves of members of the order Zingiberales. We found that the number of species used for roosting increased, that home range size increased (before: mean 0.14±SD 0.08 ha; after: 0.73±0.68 ha), and that mean association indices decreased (before: 0.95±0.10; after: 0.77±0.18) once the roosting habitat was removed. These results demonstrate that the removal of roosting resources is associated with a decrease in roost-site preferences or selectivity, an increase in mobility of individuals, and a decrease in social cohesion. These responses may reduce fitness by potentially increasing energetic expenditure, predator exposure, and a decrease in cooperative interactions. Despite these potential risks, individuals never used roost-sites other than developing furled leaves, suggesting an extreme specialization that could ultimately jeopardize the long-term persistence of this species' local populations.     

The ecological consequences of environmentally induced phenotypic changes

Population dynamics and species persistence are often mediated by species traits. Yet many important traits, like body size, can be set by resource availability and predation risk. Environmentally induced changes in resource levels or predation risk may thus have downstream ecological consequences. Here, we assess whether quantity and type of resources affect the phenotype, the population dynamics, and the susceptibility to predation of a mixotrophic protist through experiments and a model. We show that cell shape, but not size, changes with resource levels and type, and is linked to carrying capacity, thus affecting population dynamics. Also, these changes lead to differential susceptibility to predation, with direct consequences for predator‐prey dynamics. We describe important links between environmental changes, traits, population dynamics and ecological interactions, that underscore the need to further understand how trait‐mediated interactions may respond to environmental shifts in resource levels in an increasingly changing world.     

Habitat selection, facilitation, and biotic settlement cues affect distribution and performance of coral recruits in French Polynesia

Habitat selection can determine the distribution and performance of individuals if the precision with which sites are chosen corresponds with exposure to risks or resources. Contrastingly, facilitation can allow persistence of individuals arriving by chance and potentially maladapted to local abiotic conditions. For marine organisms, selection of a permanent attachment site at the end of their larval stage or the presence of a facilitator can be a critical determinant of recruitment success. In coral reef ecosystems, it is well known that settling planula larvae of reef-building corals use coarse environmental cues (i.e., light) for habitat selection. Although laboratory studies suggest that larvae can also use precise biotic cues produced by crustose coralline algae (CCA) to select attachment sites, the ecological consequences of biotic cues for corals are poorly understood in situ. In a field experiment exploring the relative importance of biotic cues and variability in habitat quality to recruitment of hard corals, pocilloporid and acroporid corals recruited more frequently to one species of CCA, Titanoderma prototypum, and significantly less so to other species of CCA; these results are consistent with laboratory assays from other studies. The provision of the biotic cue accurately predicted coral recruitment rates across habitats of varying quality. At the scale of CCA, corals attached to the “preferred” CCA experienced increased survivorship while recruits attached elsewhere had lower colony growth and survivorship. For reef-building corals, the behavioral selection of habitat using chemical cues both reduces the risk of incidental mortality and indicates the presence of a facilitator.     

Landscape scale, heterogeneity, and the viability of Serengeti grazers

Species persistence can be threatened by substantial temporal variation in food resources over time. On the other hand, spatial heterogeneity in resources at the landscape scale might allow mobile consumers to compensate for temporal variability in resource availability at the local scale. We evaluated this hypothesis, using an extensive data set on foraging, grass growth, and movement by Thomson's gazelles living on the Serengeti Plains. Here we show that modelled populations of Thomson's gazelles can only persist under Serengeti conditions in the face of observed levels of rainfall stochasticity by making adaptive movements to take advantage of ephemeral spatial distributions of food resources. More importantly, our models suggest that Thomson's gazelles in Serengeti require unrestricted access to relatively large areas of grassland (> 1600 km²) to guarantee long‐term persistence, particularly when there is positive spatial autocorrelation in resource abundance, as is the case in Serengeti. If this proves to be true for other species and/or other systems, then understanding of complex behavioural responses to spatially and temporally heterogeneous food supplies may be essential to successful conservation of grazing herbivores.     

Developmental plasticity and reduced susceptibility to natural enemies following host plant defoliation in a specialized herbivore

Host-specific phytophagous insects that are short lived and reliant on ephemeral plant tissues provide an excellent system in which to investigate the consequences of disruption in the timing of resource availability on consumer populations and their subsequent interactions with higher tropic levels. The specialist herbivore, Belonocnema treatae (Hymenoptera: Cynipidae) induces galls on only newly flushed leaves of live oak, Quercus fusiformis. In central Texas (USA) episodic defoliation of the host creates variation in the timing of resource availability and results in heterogeneous populations of B. treatae that initiate development at different times. We manipulated the timing of leaf flush in live oak via artificial defoliation to test the hypothesis that a 6- to 8-week delay in the availability of resources alters the timing of this gall former’s life cycle events, performance and survivorship on its host, and susceptibility to natural enemies. B. treatae exhibits plasticity in development time, as the interval from egg to emergence was significantly reduced when gallers oviposited into the delayed leaf flush. As a consequence, the phenologies of gall maturation and adult emergence remain synchronized in spite of variation in the timing of resource availability. Per capita gall production and gall-former performance are not significantly affected by the timing of resource availability. The timing of resource availability and natural enemies interact, however, to produce strong effects on survivorship: when exposed to natural enemies, B. treatae developing in galls initiated by delayed oviposition exhibited an order-of-magnitude increase in survivorship. Developmental plasticity allows this gall former to circumvent disruptions in resource availability, maintain synchrony of life cycle events, and results in reduced vulnerability to natural enemies following defoliation of the host plant.     

Metapopulation extinction risk is increased by environmental stochasticity and assemblage complexity

Extinction risk is a key area of investigation for contemporary ecologists and conservation biologists. Practical conservation efforts for vulnerable species can be considerably enhanced by thoroughly understanding the ecological processes that interact to determine species persistence or extinction. Theory has highlighted the importance of both extrinsic environmental factors and intrinsic demographic processes. In laboratory microcosms, single-species single-habitat patch experimental designs have been widely used to validate the theoretical prediction that environmental heterogeneity can increase extinction risk. Here, we develop on this theme by testing the effects of fluctuating resource levels in experimental multispecies metapopulations. We compare a three-species host-parasitoid assemblage that exhibits apparent competition to the individual pairwise, host-parasitoid interactions. Existing theory is broadly supported for two-species assemblages: environmental stochasticity reduces trophic interaction persistence time, while metapopulation structure increases persistence time. However, with increasing assemblage complexity, the effects of trophic interactions mask environmental impacts and persistence time is further reduced, regardless of resource renewal regime. We relate our findings to recent theory, highlighting the importance of taking into account both intrinsic and extrinsic factors, over a range of spatial scales, in order to understand resource-consumer dynamics.     

Impacts of consumer–resource interaction transitions on persistence and long‐term interaction outcomes of random ecological networks

Despite the prevalence of context‐dependent interaction transitions in ecological systems, their impacts on persistence and interaction diversity have scarcely been explored in complex ecological networks. By using multispecies bi‐directional and unidirectional consumer–resource models, representing a continuum of interaction transitions (sign change of interaction outcomes), we investigated the effects of structural interaction transitions on persistence (the fraction of remaining species) and long‐term interaction outcomes in random ecological networks. We found that high interaction strength of exploiting resources generally decreased persistence, and high strength of providing resources increased persistence when the strength of exploiting resources was low in more complex networks; also, the networks with high persistence had a high proportion of mutualistic interactions relative to antagonistic interactions present initially and over the long term. The shifting of interaction strengths shaped the long‐term interaction compositions. Meanwhile, population dynamics, especially species extinction, affected the difference between initial and long‐term interactions. Based on classical consumer–resource theory, these results establish a transitional continuum of interaction outcomes in ecological networks and imply a theoretical association among interaction transition, community persistence and interaction diversity.     

The effects of neighbors on the growth and survival of shrub seedlings following fire

Where plant species vie for limited resources, disturbances might preclude competition by releasing a flush of nutrients, or by reducing biomass and thereby diminishing the consumption of resources. However, if new seedlings colonize in clumps, they may still deplete resources within the local aggregations, which may then reduce their growth and survivorship. We investigated competition among seedlings in a burned area by examining the relationship between the performance of newly established shrub seedlings of Ceanothus impressus and (1) the proximity and (2) the identity of their near neighbors. We also investigated the relationship between neighbor proximity and the availability of water. Both survivorship and growth of C. impressus were positively associated with increasing distance to near neighbors, in a manner consistent with resource competition. The availability of water (as determined by pre-dawn xylem pressure potentials) tended to be greater when neighbors were farther away, providing evidence that water was a resource for which plants were competing. This conclusion is reinforced by the finding that the effects of neighbors were stronger in drought years, suggesting that yearly variation in the availability of an important resource (water) can affect the strength of competitive interactions. This suggests that after disturbances, when some resources are apparently abundant on a large scale, competition may be important in determining the small scale patterns of seedling growth and survival.     

Effects of partitioning allochthonous and autochthonous resources on food web stability

The flux of energetic and nutrient resources across habitat boundaries can exert major impacts on the dynamics of the recipient food web. Competition for these resources can be a key factor structuring many ecological communities. Competition theory suggests that competing species should exhibit some partitioning to minimize competitive interactions. Species should partition both in situ (autochthonous) resources and (allochthonous) resources that enter the food web from outside sources. Allochthonous resources are important sources of energy and nutrients in many low productivity systems and can significantly influence community structure. The focus of this paper is on: (i) the influence of resource partitioning on food web stability, but concurrently we examine the compound effects of; (ii) the trophic level(s) that has access to allochthonous resources; (iii) the amount of allochthonous resource input; and (iv) the strength of the consumer–resource interactions. We start with a three trophic level food chain model (resource–consumer–predator) and separate the higher two trophic levels into two trophospecies. In the model, allochthonous resources are either one type available to both consumers and predators or two distinct types, one for consumers and one for predators. The feeding preferences of the consumer and predator trophospecies were varied so that they could either be generalists or specialists on allochthonous and/or autochthonous resources. The degree of specialization influenced system persistence by altering the structure and, therefore, the indirect effects of the food web. With regard to the trophic level(s) that has access to allochthonous resources, we found that a single allochthonous resource available to both consumers and predators is more unstable than two allochthonous resources. The results demonstrate that species populating food webs that experience low to moderate allochthonous resources are more persistent. The results also support the notion that strong links destabilize food web dynamics, but that weak to moderate strength links stabilize food web dynamics. These results are consistent with the idea that the particular structure, resource availability, and relative strength of links of food webs (such as degree of specialization) can influence the stability of communities. Given that allochthonous resources are important resources in many ecosystems, we argue that the influence of such resources on species and community persistence needs to be examined more thoroughly to provide a clearer understanding of food web dynamics.     

Corridors and barriers in biodiversity conservation: a novel resource-based habitat perspective for butterflies

Habitat loss and fragmentation, exacerbated by projected climate change, present the greatest threats to preservation of global biodiversity. As increasing habitat fragmentation and isolation of residual fragments exceeds the dispersal capacity of species, there is the growing need to address connectivity to maintain diversity. Traditionally, habitat corridors have been proposed as a solution. But, the concept of corridors (barriers) is poorly understood; typically they are defined as linear habitats linking up habitat patchwork, and are advocated without a detailed understanding of the elements making up species’ habitats and the cost-effectiveness of alternative solutions. Yet, landscapes comprise an enormous range of ‘linear’ structures that can function in different ways to promote species’ persistence and diversity. In this review, a functional definition of corridor (barrier) is developed to give prominence to connectivity as opposed to ad hoc structures purported to advance connectivity. In developing the concept, urgency to accommodate environmental changes compels a growing emphasis on organism diversity rather than a preoccupation with single species conservation. The review, in focusing on butterflies to address the issue of corridors for patchwork connectivity, draws attention to fundamental divisions among organisms in any taxon: generalists and specialists. Both groups benefit from large patches as these necessarily house species with specialist resources as well as generalists with very different resource types. But, generalists and specialists require very different solutions for connectivity, from short-range habitat corridors and gateways for specialists to habitat and resource stepping stones (nodes, surfaces) for generalists. Connectivity over extensive areas is most critical for moderate generalists and their conservation requires emphasis being placed on space–time resource heterogeneity; landscape features, of whatever dimensionality and structure, provide a vital framework for developing the variety of suitable conditions and resources for enhancing their diversity.     

Factors underlying the late seasonal appearance of the lepidopterous leaf-mining guild on oak

Abstract. 1. Most lepidopterous leaf mining species found on the oak Quercus robur in Britain develop in summer. At this time of year, externally feeding caterpillars remove little leaf area since most of these free living folivores are predominantly spring feeders.
2. I forced Phyllonorycter harrisella (L.) miners to oviposit in spring, then exposed developing larvae to a wide range of leaf damage levels.
3. Leaf miner survivorship and mean female pupal weight were significantly greater in the experimental spring generation on undamaged oak leaves, this being when oak foliage is of the highest nutritional quality.
4. Leaf miner survivorship in all generations is correlated with natural leaf damage levels. Experimental leaf damage also reduces miner survivorship.
5. Leaf damage reduces miner survivorship by increasing the probability of larval death due to wound induced responses.
6. The role of asymmetrical competitive interactions between caterpillars and leaf-miners in determining the late seasonal appearance of miners is discussed. It is suggested that wound induced responses may play an important part in the structuring of phytophagous insect communities.     

Host–parasitoid persistence over variable spatio‐temporally susceptible habitats: bottom–up effects of ephemeral resources

We experimentally and theoretically investigated the persistence of hosts and parasitoids interacting in a metapopulation structure consisting of ephemeral local patches (MELPs). We used a host–parasitoid system consisting of necrophagous Diptera species and their pupal parasitoids. The basal resources used by the host species were assumed to be ephemeral, supporting only one generation of individuals before completely disappearing from the environment. We experimentally measured the host–parasitoid persistence and the effects of local demographic processes in two scenarios: 1) constant occurrence of basal resources at a single site (no dispersion or colonization of other sites) and 2) variable occurrence of basal resources between two sites (colonization of a new patch requiring species dispersal). The experimental setup and findings were then formalized into a mathematical model describing the interaction dynamics in a MELP structure. We evaluated the contribution of several factors to the host–parasitoid coexistence, such as resource allocation probability (probability of resource appearance in a site), variation in resource size and number of sites available to receive resources in the MELP. We found that demographic fluctuations and environmental stochasticity affected the density of migrants, patch habitat connectivity, persistence and spatial distribution of interacting species.     

Temporal resource partitioning mitigates interspecific competition and promotes coexistence among insect parasites

A key to understanding life's great diversity is discerning how competing organisms divide limiting resources to coexist in diverse communities. While temporal resource partitioning has long been hypothesized to reduce the negative effects of interspecific competition, empirical evidence suggests that time may not often be an axis along which animal species routinely subdivide resources. Here, we present evidence to the contrary in the world's most biodiverse group of animals: insect parasites (parasitoids). Specifically, we conducted a meta-analysis of 64 studies from 41 publications to determine if temporal resource partitioning via variation in the timing of a key life-history trait, egg deposition (oviposition), mitigates interspecific competition between species pairs sharing the same insect host. When competing species were manipulated to oviposit at (or near) the same time in or on a single host in the laboratory, competition was common, and one species was typically inherently superior (i.e. survived to adulthood a greater proportion of the time). In most cases, however, the inferior competitor could gain a survivorship advantage by ovipositing earlier (or in a smaller number of cases later) into shared hosts. Moreover, this positive (or in a few cases negative) priority advantage gained by the inferior competitor increased as the interval between oviposition times became greater. The results from manipulative experiments were also correlated with patterns of life-history timing and demography in nature: the more inherently competitively inferior a species was in the laboratory, the greater the interval between oviposition times of taxa in co-occurring populations. Additionally, the larger the interval between oviposition times of competing taxa, the more abundant the inferior species was in populations where competitors were known to coexist. Overall, our findings suggest that temporal resource partitioning via variation in oviposition timing may help to facilitate species coexistence and structures diverse insect communities by altering demographic measures of species success. We argue that the lack of evidence for a more prominent role of temporal resource partitioning in promoting species coexistence may reflect taxonomic differences, with a bias towards larger-sized animals. For smaller species like parasitic insects that are specialized to attack one or a group of closely related hosts, have short adult lifespans and discrete generation times, compete directly for limited resources in small, closed arenas and have life histories constrained by host phenology, temporal resource subdivision via variation in life history may play a critical role in allowing species to coexist by alleviating the negative effects of interspecific competition.     

Resource selection and its implications for wide-ranging mammals of the brazilian cerrado

Conserving animals beyond protected areas is critical because even the largest reserves may be too small to maintain viable populations for many wide-ranging species. Identification of landscape features that will promote persistence of a diverse array of species is a high priority, particularly, for protected areas that reside in regions of otherwise extensive habitat loss. This is the case for Emas National Park, a small but important protected area located in the Brazilian Cerrado, the world's most biologically diverse savanna. Emas Park is a large-mammal global conservation priority area but is too small to protect wide-ranging mammals for the long-term and conserving these populations will depend on the landscape surrounding the park. We employed novel, noninvasive methods to determine the relative importance of resources found within the park, as well as identify landscape features that promote persistence of wide-ranging mammals outside reserve borders. We used scat detection dogs to survey for five large mammals of conservation concern: giant armadillo (Priodontes maximus), giant anteater (Myrmecophaga tridactyla), maned wolf (Chrysocyon brachyurus), jaguar (Panthera onca), and puma (Puma concolor). We estimated resource selection probability functions for each species from 1,572 scat locations and 434 giant armadillo burrow locations. Results indicate that giant armadillos and jaguars are highly selective of natural habitats, which makes both species sensitive to landscape change from agricultural development. Due to the high amount of such development outside of the Emas Park boundary, the park provides rare resource conditions that are particularly important for these two species. We also reveal that both woodland and forest vegetation remnants enable use of the agricultural landscape as a whole for maned wolves, pumas, and giant anteaters. We identify those features and their landscape compositions that should be prioritized for conservation, arguing that a multi-faceted approach is required to protect these species.     

Resource discovery in ant communities: do food type and quantity matter?

1. Omnivorous woodland ant species trade off between the ability to find and behaviourally control food resources. Dominant species can limit the ability of subordinates to harvest certain food items. However, subordinate species, by being faster discoverers, could gain access to such food items by arriving at them first. 2. In this study, we tested the hypothesis that resource‐directed discovery occurs in ant communities and that good discoverers preferentially discover high value resources. We did this by measuring time to discovery and the number of discoveries of high and low levels of two resource types, crickets and honey, for species occurring in Texas and Arizona woodland ant communities. 3. Ants discovered resources roughly 10 times faster in Texas than in Arizona. They discovered crickets more rapidly than honey in both communities, but there was no difference in the discovery of different resource levels. We also found that species were not biased in their discovery of different resource types or levels. 4. These results provide indirect evidence that discovery is directed by resource stimuli but that such directedness does not impact interspecific exploitative competition.