Pattern of seasonal variation in rates of predation between spider families is temporally stable in a food web with widespread intraguild predation.

Intraguild predation (IGP)-predation between generalist predators (IGPredator and IGPrey) that potentially compete for a shared prey resource-is a common interaction module in terrestrial food webs. Understanding temporal variation in webs with widespread IGP is relevant to testing food web theory. We investigated temporal constancy in the structure of such a system: the spider-focused food web of the forest floor. Multiplex PCR was used to detect prey DNA in 3,300 adult spiders collected from the floor of a deciduous forest during spring, summer, and fall over four years. Because only spiders were defined as consumers, the web was tripartite, with 11 consumer nodes (spider families) and 22 resource nodes: 11 non-spider arthropod taxa (order- or family-level) and the 11 spider families. Most (99%) spider-spider predation was on spider IGPrey, and ~90% of these interactions were restricted to spider families within the same broadly defined foraging mode (cursorial or web-spinning spiders). Bootstrapped-derived confidence intervals (BCI's) for two indices of web structure, restricted connectance and interaction evenness, overlapped broadly across years and seasons. A third index, % IGPrey (% IGPrey among all prey of spiders), was similar across years (~50%) but varied seasonally, with a summer rate (65%) ~1.8x higher than spring and fall. This seasonal pattern was consistent across years. Our results suggest that extensive spider predation on spider IGPrey that exhibits consistent seasonal variation in frequency, and that occurs primarily within two broadly defined spider-spider interaction pathways, must be incorporated into models of the dynamics of forest-floor food webs.

Effects of enhanced productivity of resources shared by predators in a food-web module: Comparing results of a field experiment to predictions of mathematical models of intra-guild predation.

We compared the response to resource enhancement of a simple empirical model of intra-guild predation (IGP) to the predictions of published, simple mathematical models of asymmetric IGP (a generalist IG Predator that feeds both on a specialist IG Prey and a Resource that it shares with the IG Prey). The empirical model was a food-web module created by pooling species abundances across many families in a speciose community of soil micro-arthropods into three categories: IG Predator (large predatory mites), IG Prey (small predatory mites), and a shared Resource (fungivorous mites and springtails). By pooling abundances of species belonging to broadly defined functional groups, we tested the hypothesis that IGP is a dominant organizing principle in this community. Simple mathematical models of asymmetric IGP predict that increased input of nutrients and energy to the shared Resource will increase the equilibrium density of Resource and IG Predator, but will decrease that of IG Prey. In a field experiment, we observed how the three categories of the empirical model responded to two rates of addition of artificial detritus, which enhanced the food of fungivores, the Resource of the IGP module. By the experiment's end, fungivore densities had increased ~1.5× (ratio of pooled fungivore densities in the higher-input treatment to plots with no addition of detritus), and densities of IG Predators had increased ~4×. Contrary to the prediction of mathematical models, IG Prey had not decreased, but instead had increased ~1.5×. We discuss possible reasons for the failure of the empirical model to agree with IGP theory. We then explore analogies between the behavior of the empirical model and another mathematical model of trophic interactions as one way to gain insights into the trophic connections in this community. We also propose one way forward for reporting comparisons of simple empirical and mathematical models.

Rodent suppression of seedling establishment in tropical pasture.

Grasses are recognized as a critical regeneration barrier in tropical pastures, yet the effects of rodents and rodent-grass interactions are not well understood. As selective foragers, rodents could shape tree communities, moderating biodiversity in regenerating tropical landscapes. We utilized a fully crossed two-way factorial design to examine the effect that grasses, rodents, and their interaction had on tree seedling establishment in pasture habitat. We followed two separate tree cohorts for 1 year each within the experimental framework. Multiple cohorts were used to better represent successional tree species variation and responses. Trees species were characterized by a gradient of seed masses and as pioneer or persistent successional type. Both cohort seedlings were altered when rodents were present compared to control treatments. In Cohort 1, rodents adversely affected seedlings of persistent tree species only in the absence of grass. In Cohort 2, seedlings of persistent tree species were decimated by rodents in the absence or presence of grass. In both cohorts, seedlings of persistent species established better in grass treatments, while seedlings of pioneer tree species were strongly suppressed. Tree species seed mass positively correlated with seedling establishment across all treatments except no grass-rodent treatments. Strong suppression of tree seedlings by rodents (Sigmodon toltecus) is a novel result in tropical land recently released from agriculture. One implication is that selective foraging by rodents on large-seeded persistent tree species may be facilitated by the removal of grass. Another implication is that temporary rodent control in pastures may permit higher establishment of deep-forest persistent species.

Field exclusion of large soil predators impacts lower trophic levels and decreases leaf-litter decomposition in dry forests.

Shifts in densities of apex predators may indirectly affect fundamental ecosystem processes, such as decomposition, by altering patterns of cascading effects propagating through lower trophic levels. These top-down effects may interact with anthropogenic impacts, such as climate change, in largely unknown ways. We investigated how changes in densities of large predatory arthropods in forest leaf-litter communities altered lower trophic levels and litter decomposition. We conducted our experiment in soil communities that had experienced different levels of long-term average precipitation. We hypothesized that altering abundances of apex predators would have stronger effects on soil communities inhabiting dry forests, due to lower secondary productivity and greater resource overexploitation by lower trophic levels compared to wet forests. We experimentally manipulated abundances of the largest arthropod predators (apex predators) in field mesocosms replicated in the leaf-litter community of Iberian beech forests that differed in long-term mean annual precipitation by 25% (three dry forests with MAP < 1,250 mm and four wet forests with MAP > 1,400 mm). After one year, we assessed abundances of soil fauna in lower trophic levels and indirect impacts on leaf-litter decomposition using litter of understorey hazel, Corylus avellana. Reducing densities of large predators had a consistently negative effect on final abundances of the different trophic groups and several taxa within each group. Moreover, large predatory arthropods strongly impacted litter decomposition, and their effect interacted with the long-term annual rainfall experienced by the soil community. In the dry forests, a 50% reduction in the densities of apex predators was associated with a 50% reduction in decomposition. In wet forests, the same reduction in densities of apex soil predators did not alter the rate of litter decomposition. Our results suggest that predators may facilitate lower trophic levels by indirectly reducing competition and resource overexploitation, cascading effects that may be more pronounced in drier forests where conditions have selected for greater competitive ability and more rapid resource utilization. These findings thus provide insights into the functioning of soil invertebrate communities and their role in decomposition, as well as potential consequences of soil community responses to climate change.

Plant invader alters soil food web via changes to fungal resources.

While aboveground impacts of invasive plants are well documented, their influence on soil food webs remains less understood. Previous research has revealed that bottom-up forces are widespread in soil food webs of woodlands. Thus, an invasive plant that negatively impacts the base of the food web will likely decrease primary consumers as well as their predators. We examined how a North American plant invader, garlic mustard (Alliaria petiolata), affects arthropod primary (springtails and oribatid mites) and secondary (predaceous mites) consumers of the soil food web via changes to fungal resources. We measured the abundances of plants, soil fungi, fungivores, and predators in garlic mustard-invaded and uninvaded 1-m2 plots in five Midwestern USA woodlands. We then conducted a mesocosm (0.25-m2 plots) experiment to tease apart the direct and indirect effects of garlic mustard by manipulating plant identity (garlic mustard vs. native plant), soil history (invaded vs. uninvaded), and fungicide application (fungicide vs. no fungicide). Our first study revealed that plots without garlic mustard had 2.8 and 1.4 × more fungi and fungivores, respectively. Predator densities did not differ. Fungal composition and structural equation modeling (SEM) revealed the garlic mustard effects on fungivores were correlated with fungal declines. The mesocosm experiment confirmed that the impacts were indirect, as fungicide plots harbored similar fungivore densities, whereas fungivore densities differed according to plant identity and soil history in the fungicide-free plots. Our results reveal that by altering soil fungal abundance, an invasive plant can indirectly affect primary consumers in soil food webs, but this indirect effect does not influence predators.

Impacts of rainfall extremes predicted by climate-change models on major trophic groups in the leaf litter arthropod community.

Arthropods in the leaf litter layer of forest soils influence ecosystem processes such as decomposition. Climate-change models predict both increases and decreases in average rainfall. Increased drought may have greater impacts on the litter arthropod community. In addition to affecting survival or behaviour of desiccation-sensitive species, lower rainfall may indirectly lower abundances of consumers that graze drought-stressed fungi, with repercussions for higher trophic levels. We tested the hypothesis that trophic structure will differ between the two rainfall scenarios. In particular, we hypothesized that densities of several broadly defined trophic groupings of arthropods would be lower under reduced rainfall. To test this hypothesis, we used sprinklers to impose two rainfall treatments during three growing seasons in roofed, fenced 14-m2 plots and documented changes in abundance from initial, pre-treatment densities of 39 arthropod taxa. Experimental plots were subjected to either LOW (fortnightly) or HIGH (weekly) average rainfall based upon climate models and the previous 100 years of regional weekly averages. Unroofed open plots, our reference treatment (REF), experienced higher than average rainfall during the experiment. The two rainfall extremes produced clear negative effects of lowered rainfall on major trophic groups. Broad categories of fungivores, detritivores and predators were more abundant in HIGH than LOW plots by the final year. Springtails (Collembola), which graze fungal hyphae, were 3× more abundant in the HIGH rainfall treatment. Taxa of larger-bodied fungivores and detritivores, spiders (Araneae), and non-spider predators were 2× more abundant under HIGH rainfall. Densities of mites (Acari), which include fungivores, detritivores and predators, were 1.5× greater in HIGH rainfall plots. Abundances and community structure of arthropods were similar in REF and experimental plots, showing that effects of rainfall uncovered in the experiment are applicable to nature. This pattern suggests that changes in rainfall will alter bottom-up control processes in a critical detritus-based food web of deciduous forests. Our results, in conjunction with other findings on the impact of desiccation on arthropods and fungal growth, suggest that drier conditions will depress densities of fungal consumers, causing declines in higher trophic levels, with possible impacts on soil processes and the larger forest food web.

The role of plant-mycorrhizal mutualisms in deterring plant invasions: Insights from an individual-based model.

Understanding the factors that determine invasion success for non-native plants is crucial for maintaining global biodiversity and ecosystem functioning. One hypothesized mechanism by which many exotic plants can become invasive is through the disruption of key plant-mycorrhizal mutualisms, yet few studies have investigated how these disruptions can lead to invader success. We present an individual-based model to examine how mutualism strengths between a native plant (Impatiens capensis) and mycorrhizal fungus can influence invasion success for a widespread plant invader, Alliaria petiolata (garlic mustard). Two questions were investigated as follows: (a) How does the strength of the mutualism between the native I. capensis and a mycorrhizal fungus affect resistance (i.e., native plant maintaining >60% of final equilibrium plant density) to garlic mustard invasion? (b) Is there a non-linear relationship between initial garlic mustard density and invasiveness (i.e., garlic mustard representing >60% of final equilibrium plant density)? Our findings indicate that either low (i.e., facultative) or high (i.e., obligate) mutualism strengths between the native plant and mycorrhizal fungus were more likely to lead to garlic mustard invasiveness than intermediate levels, which resulted in higher resistance to garlic mustard invasion. Intermediate mutualism strengths allowed I. capensis to take advantage of increased fitness when the fungus was present but remained competitive enough to sustain high numbers without the fungus. Though strong mutualisms had the highest fitness without the invader, they proved most susceptible to invasion because the loss of the mycorrhizal fungus resulted in a reproductive output too low to compete with garlic mustard. Weak mutualisms were more competitive than strong mutualisms but still led to garlic mustard invasion. Furthermore, we found that under intermediate mutualism strengths, the initial density of garlic mustard (as a proxy for different levels of plant invasion) did not influence its invasion success, as high initial densities of garlic mustard did not lead to it becoming dominant. Our results indicate that plants that form weak or strong mutualisms with mycorrhizal fungi are most vulnerable to invasion, whereas intermediate mutualisms provide the highest resistance to an allelopathic invader.

Long-term resource addition to a detrital food web yields a pattern of responses more complex than pervasive bottom-up control.

BACKGROUND: Theory predicts strong bottom-up control in detritus-based food webs, yet field experiments with detritus-based terrestrial systems have uncovered contradictory evidence regarding the strength and pervasiveness of bottom-up control processes. Two factors likely leading to contradictory results are experiment duration, which influences exposure to temporal variation in abiotic factors such as rainfall and affects the likelihood of detecting approach to a new equilibrium; and openness of the experimental units to immigration and emigration. To investigate the contribution of these two factors, we conducted a long-term experiment with open and fenced plots in the forest that was the site of an earlier, short-term experiment (3.5 months) with open plots (Chen & Wise, 1999) that produced evidence of strong bottom-up control for 14 taxonomic groupings of primary consumers of fungi and detritus (microbi-detritivores) and their predators. METHODS: We added artificial high-quality detritus to ten 2 × 2-m forest-floor plots at bi-weekly intervals from April through September in three consecutive years (Supplemented treatment). Ten comparable Ambient plots were controls. Half of the Supplemented and Ambient plots were enclosed by metal fencing. RESULTS: Arthropod community structure (based upon 18 response variables) diverged over time between Supplemented and Ambient treatments, with no effect of Fencing on the multivariate response pattern. Fencing possibly influenced only ca. 30% of the subsequent univariate analyses. Multi- and univariate analyses revealed bottom-up control during Year 1 of some, but not all, microbi-detritivores and predators. During the following two years the pattern of responses became more complex than that observed by Chen & Wise (1999). Some taxa showed consistent bottom-up control whereas others did not. Variation across years could not be explained completely by differences in rainfall because some taxa exhibited negative, not positive, responses to detrital supplementation. DISCUSSION: Our 3-year experiment did not confirm the conclusion of strong, pervasive bottom-up control of both microbi-detritivores and predators reported by Chen & Wise (1999). Our longer-term experiment revealed a more complex pattern of responses, a pattern much closer to the range of outcomes reported in the literature for many short-term experiments. Much of the variation in responses across studies likely reflects variation in abiotic and biotic factors and the quality of added detritus. Nevertheless, it is also possible that long-term resource enhancement can drive a community towards a new equilibrium state that differs from what would have been predicted from the initial short-term responses exhibited by primary and secondary consumers.

Substantial Mortality of Cabbage Looper (Lepidoptera: Noctuidae) From Predators in Urban Agriculture Is not Influenced by Scale of Production or Variation in Local and Landscape-Level Factors.

As Midwestern (United States) cities experience population decline, there is growing interest in converting underutilized vacant spaces to agricultural production. Urban agriculture varies in area and scope, yet most growers use similar cultivation practices such as avoiding chemical control of crop pests. For community gardens and farms that sell produce commercially, effective pest suppression by natural enemies is important for both societal, economic, and marketing reasons. To gauge the amount of prey suppression at 28 urban food-production sites, we measured removal of sentinel eggs and larvae of the cabbage looper Trichoplusia ni (Hubner), a caterpillar pest that defoliates Brassica. We investigated how landscape and local factors, such as scale of production, influence cabbage looper mortality caused by predators. Predators removed 50% of eggs and 25% of larvae over a 3-d period. Landscape factors did not predict mortality rates, and the amount of loss and damage to sentinel prey were similar across sites that differed in scale (residential gardens, community gardens, and farms). To confirm that removal of sentinel items was likely caused by natural enemies, we set up a laboratory assay that measured predation of cabbage looper eggs and larvae by several predators occurring in urban gardens. Lady beetles caused the highest mortality rates, suggesting their potential value for biocontrol; spiders and pirate bugs also consumed both eggs and larvae at high rates. Our results suggest that urban growers benefit from high consumption rates of cabbage looper eggs and larvae by arthropod predators.

Invasive plants have different effects on trophic structure of green and brown food webs in terrestrial ecosystems: a meta-analysis.

Although invasive plants are a major source of terrestrial ecosystem degradation worldwide, it remains unclear which trophic levels above the base of the food web are most vulnerable to plant invasions. We performed a meta-analysis of 38 independent studies from 32 papers to examine how invasive plants alter major groupings of primary and secondary consumers in three globally distributed ecosystems: wetlands, woodlands and grasslands. Within each ecosystem we examined if green (grazing) food webs are more sensitive to plant invasions compared to brown (detrital) food webs. Invasive plants have strong negative effects on primary consumers (detritivores, bacterivores, fungivores, and/or herbivores) in woodlands and wetlands, which become less abundant in both green and brown food webs in woodlands and green webs in wetlands. Plant invasions increased abundances of secondary consumers (predators and/or parasitoids) only in woodland brown food webs and green webs in wetlands. Effects of invasive plants on grazing and detrital food webs clearly differed between ecosystems. Overall, invasive plants had the most pronounced effects on the trophic structure of wetlands and woodlands, but caused no detectable changes to grassland trophic structure.

Modeling the impacts of life-history traits, canopy gaps, and establishment location on woodland shrub invasions.

We used an individual-based model to identify how localized patterns of woodland invasions by exotic shrubs are likely influenced by (1) observed variation in age at first reproduction and fecundity, (2) hypothesized effects of canopy gaps on these life-history traits and dispersal, and (3) initial establishment location. Rates of spread accelerated nearly twofold as age at first reproduction decreased from eight to three years or fecundity increased from 3 to 20 offspring per year, illustrating the need to better understand the factors that influence these life-history traits. Canopy gaps facilitated spread by influencing these life-history traits, but not through their effects on dispersal. Invasions starting at the woodland center spread more rapidly than do those starting along the woodland edge. These findings suggest that managers should not only prioritize the removal of shrubs that reproduce the earliest or produce the most offspring, but they should also focus on the invasions in woodlands with high canopy openness and/or that are located in woodland interiors. Investigated factors also affected other invasion characteristics, often in surprising ways. For example, those changes in age at first reproduction and fecundity that increased the rate of spread produced nonparallel patterns of change in the proportions of invasion reproducing, whether or not invasions exhibited clumped or scattered spatial arrangements, and invasional lag. Additionally, canopy gaps influenced these characteristics by increasing fecundity, but not by decreasing age at first reproduction or altering dispersal, suggesting that canopy gaps affect local patterns of exotic-shrub invasions primarily through their positive effects on fruit production.

Coexistence and community structure in a consumer resource model with implicit stoichiometry.

We combine stoichiometry theory and optimal foraging theory into the MacArthur consumer-resource model. This generates predictions for diet choice, coexistence, and community structure of heterotroph communities. Tradeoffs in consumer resource-garnering traits influence community outcomes. With scarce resources, consumers forage opportunistically for complementary resources and may coexist via tradeoffs in resource encounter rates. In contrast to single currency models, stoichiometry permits multiple equilibria. These alternative stable states occur when tradeoffs in resource encounter rates are stronger than tradeoffs in elemental conversion efficiencies. With abundant resources consumers exhibit partially selective diets for essential resources and may coexist via tradeoffs in elemental conversion efficiencies. These results differ from single currency models, where adaptive diet selection is either opportunistic or selective. Interestingly, communities composed of efficient consumers share many of the same properties as communities based on substitutable resources. However, communities composed of relatively inefficient consumers behave similarly to plant communities as characterized by Tilman's consumer resource theory. The results of our model indicate that the effects of stoichiometry theory on community ecology are dependent upon both consumer foraging behavior and the nature of resource garnering tradeoffs.

Sexual cannibalism: high incidence in a natural population with benefits to females.

BACKGROUND: Sexual cannibalism may be a form of extreme sexual conflict in which females benefit more from feeding on males than mating with them, and males avoid aggressive, cannibalistic females in order to increase net fitness. A thorough understanding of the adaptive significance of sexual cannibalism is hindered by our ignorance of its prevalence in nature. Furthermore, there are serious doubts about the food value of males, probably because most studies that attempt to document benefits of sexual cannibalism to the female have been conducted in the laboratory with non-natural alternative prey. Thus, to understand more fully the ecology and evolution of sexual cannibalism, field experiments are needed to document the prevalence of sexual cannibalism and its benefits to females. METHODOLOGY/PRINCIPAL FINDINGS: We conducted field experiments with the Mediterranean tarantula (Lycosa tarantula), a burrowing wolf spider, to address these issues. At natural rates of encounter with males, approximately a third of L. tarantula females cannibalized the male. The rate of sexual cannibalism increased with male availability, and females were more likely to kill and consume an approaching male if they had previously mated with another male. We show that females benefit from feeding on a male by breeding earlier, producing 30% more offspring per egg sac, and producing progeny of higher body condition. Offspring of sexually cannibalistic females dispersed earlier and were larger later in the season than spiderlings of non-cannibalistic females. CONCLUSIONS/SIGNIFICANCE: In nature a substantial fraction of female L. tarantula kill and consume approaching males instead of mating with them. This behaviour is more likely to occur if the female has mated previously. Cannibalistic females have higher rates of reproduction, and produce higher-quality offspring, than non-cannibalistic females. Our findings further suggest that female L. tarantula are nutrient-limited in nature and that males are high-quality prey. The results of these field experiments support the hypothesis that sexual cannibalism is adaptive to females.

Small-scale spatial pattern of web-building spiders (Araneae) in alfalfa: relationship to disturbance from cutting, prey availability, and intraguild interactions.

Understanding the development of spatial patterns in generalist predators will improve our ability to incorporate them into biological control programs. We studied the small-scale spatial patterns of spider webs in alfalfa by analyzing the relationship between web locations over distances ranging from 4 to 66 cm. Using a coordinate-based spatial statistic (O-ring) and assuming a heterogeneous distribution of suitable web sites, we analyzed the impact of cutting and changes in spider abundance on web distribution. We analyzed the influence of small-scale variation in prey availability by comparing web distributions to the pattern of sticky-trap captures of Aphididae and Diptera described by a count-based spatial statistic (SADIE). Cutting of alfalfa reduced the overall density of web-building spiders but had no immediate impact on the spatial distribution of their webs. Availability of aphids was highest before the alfalfa was cut and was clumped at a scale of 66 cm. Spider webs, however, were not clumped at any scale or date. In contrast, webs were regularly distributed at smaller distances (<20 cm) immediately before and after cutting. Because cursorial and web-building spiders were most active during this period, we hypothesize that the development of small-scale regularity in web locations was driven by intraguild interactions. Our results suggest that intraguild interactions contribute to the development of small-scale spatial patterns of spider webs in alfalfa. Variation in prey availability may have more of an influence on web distribution in crops with a different vegetation structure or if patterns are studied at larger spatial scales.

Direct and indirect effects of ants on a forest-floor food web.

Interactions among predators that prey on each other and are potential competitors for shared prey (intraguild [IG] predators) are widespread in terrestrial ecosystems and have the potential to strongly influence the dynamics of terrestrial food webs. Ants and spiders are abundant and ubiquitous terrestrial IG predators, yet the strength and consequences of interactions between them are largely unknown. In the leaf-litter food web of a deciduous forest in Kentucky (USA), we tested the direct and indirect effects of ants on spiders and a category of shared prey (Collembola) by experimentally subsidizing ants in open plots in two field experiments. In the first experiment, ant activity was increased, and the density of ants in the litter was doubled, by placing carbohydrate and protein baits in the center of each plot. Gnaphosa spiders were almost twice as abundant and Schizocosa spiders were half as abundant in baited plots relative to controls. There were more tomocerid Collembola in baited plots, suggesting possible indirect effects on Collembola caused by ant-spider interactions. The second experiment, in which screening of two mesh sizes selectively excluded large and small worker ants from a sugar bait, revealed that the large ants, primarily Camponotus, could alone induce similar effects on spiders. Gnaphosa biomass density was almost twice as high in the plots where large ants were more active, whereas Schizocosa biomass density was reduced by half in these plots. Although tomocerid densities did not differ between treatments, tomocerid numbers were negatively correlated with the activity of Formica, another large ant species. Path analysis failed to support the hypothesis that the ant Camponotus indirectly affected tomocerid Collembola through effects on densities of spiders. However, path analysis also revealed other indirect effects of Camponotus affecting tomocerids. These results illustrate the complexity of interactions between and within two major IG predator groups with disparate predatory behaviors, complexities that will have consequences for functioning of the forest-floor food web.

Predicted climate change alters the indirect effect of predators on an ecosystem process.

Changes in rainfall predicted to occur with global climate change will likely alter rates of leaf-litter decomposition through direct effects on primary decomposers. In a field experiment replicated at two sites, we show that altered rainfall may also change how cascading trophic interactions initiated by arthropod predators in the leaf litter indirectly influence litter decomposition. On the drier site there was no interaction between rainfall and the indirect effect of predators on decomposition. In contrast, on the moister site spiders accelerated the disappearance rate of deciduous leaf litter under low rainfall, but had no, or possibly a negative, indirect effect under high rainfall. Thus, changes resulting from the more intense hydrological cycle expected to occur with climate change will likely influence how predators indirectly affect an essential ecosystem process.

Using stable isotopes to reveal shifts in prey consumption by generalist predators.

The effectiveness of generalist predators in biological control may be diminished if increased availability of alternative prey causes individual predators to decrease their consumption of crop pests. Farming practices that enhance densities of microbidetritivores in the detrital food web can lead to increased densities of generalist predators that feed on pest species. The ability to predict the net biocontrol impact of increased predator densities depends upon knowing the extent to which individual predators may shift to detrital prey and feed less on crop pests when prey of the detritus-based food web are more abundant. We addressed this question by comparing ratios of stable isotopes of carbon (delta13C) and nitrogen (delta15N) in generalist ground predators and two types of prey (crop pests and microbidetritivores) in replicated 8 x 8 m cucurbit gardens subjected to one of two treatments: a detrital subsidy or no addition of detritus (control). Small sheet-web spiders (Linyphiidae) and small wolf spiders (Lycosidae) had delta13C values similar to those of Collembola in both the detrital and control treatments, indicating that small spiders belong primarily to the detrital food web. In control plots the larger generalist predators had delta13C values similar to those of the major insect pests, consistent with their known effectiveness as biocontrol agents. Adding detritus may have caused delta13C of one species of large wolf spider to shift toward that of the microbi-detritivores, although evidence is equivocal. In contrast, another large wolf spider displayed no shift in delta13C in the detrital treatment. Thus, stable isotopes revealed which generalist predators will likely continue to feed on pest species in the presence of greater densities of alternative prey.

Cannibalism, food limitation, intraspecific competition, and the regulation of spider populations.

Cannibalism among generalist predators has implications for the dynamics of terrestrial food webs. Spiders are common, ubiquitous arthropod generalist predators in most natural and managed terrestrial ecosystems. Thus, the relationship of spider cannibalism to food limitation, competition, and population regulation has direct bearing on basic ecological theory and applications such as biological control. This review first briefly treats the different types of spider cannibalism and then focuses in more depth on evidence relating cannibalism to population dynamics and food web interactions to address the following questions: Is cannibalism in spiders a foraging strategy that helps to overcome the effects of a limited supply of calories and/or nutrients? Does cannibalism in spiders reduce competition for prey? Is cannibalism a significant density-dependent factor in spider population dynamics? Does cannibalism dampen spider-initiated trophic cascades?

Climbing to reach females: Romeo should be small.

The race for reaching mates by the time they are receptive, or sexual selection by scramble competition, has received little attention. We argue that smaller males are favored in species in which the male must climb to reach females located in high habitat patches. This new explanation we term the "gravity hypothesis" of sexual size dimorphism (SSD). We show that a simple biomechanical model of animal movement predicts that: (1) selection should favor a comparatively smaller size in the searching sex when searching involves climbing; and (2) this effect should be stronger in larger species than in smaller species. In reaching high habitats, smaller, faster searchers will be favored either through sexual selection by scramble competition and/or by escaping predation easier by running faster on vertical surfaces. Different spider species are found at a wide range of heights. We compiled a dataset of spider taxa and arranged their habitats according to four height categories, ranked from soil surface to trees. We show that, after controlling for phylogeny, both predictions of the gravity hypothesis of SSD are met. Thus, it appears that the constraint imposed by gravity on climbing males is a selective factor in determining male dwarfism.

Prey availability limits fecundity and influences the movement pattern of female fishing spiders.

We conducted a food supplementation field experiment to test two hypotheses: (1) fecundity of the fishing spider Dolomedes triton is limited by a shortage of prey, and (2) the increased movement of adult female D. triton exhibited upon maturation is a foraging adaptation to lessen the impact of food limitation on egg production. Free ranging, uniquely marked adult female fishing spiders were assigned either to a food-supplemented group that received crickets in addition to their natural diet, or to a control group. Juvenile female spiders were also marked and their movement patterns were recorded, but juveniles were not offered supplemental food. Food-supplemented adult females gained weight at a faster rate and hatched more than twice as many offspring as control females. Adult females in the control group moved greater distances per day than did juvenile females. Supplemented adult females moved shorter distances per day than control females, and the movement pattern of fed adults did not differ from that of juveniles. These results support the hypotheses that adult female D. triton are food limited, and that the increased movement of adult females is a switch in foraging behavior that occurs during the reproductive period. Our finding that natural prey shortages limit egg production contrasts with laboratory-based studies of food limitation in the genus Dolomedes, and contradicts a basic assumption of a recent hypothesis that sexual cannibalism in Dolomedes is non-adaptive. These discrepancies highlight the importance of insights gained from field experiments with natural populations.