鼎湖山锥栗种子特征及其对动物取食格局的影响

通过对鼎湖山自然保护区锥栗种子特征的分析及进行种子取食实验，研究探讨了锥栗种子特征对动物取食格局的影响。结果表明，锥栗作为鼎湖山季风常绿阔叶林中的重要建群种之一，其更新困难的主要原因就在于种子扩散前后面临巨大的取食压力。动物对种子取食偏爱同种子大小呈现出一定的相关性，随着种子尺寸增加，取食率有线性增加的趋势。但是，锥栗应对动物取食并非完全被动的，它也发展了一些特有的生活史策略。锥栗种子的颜色与环境凋落物及土壤色相似，可以减少被取食者发现的机率。更重要的是，锥栗种子带刺的总苞有良好的保护作用。具总苞的种子取食的强度，显著低于无总苞的种子（P〈0．01）。     

Opposite shifts in size at metamorphosis in response to larval and metamorph predators

Predation risk can cause organisms to alter the timing of life history switch points. Theory suggests that increased risk in an early life stage should select for switching earlier and smaller, while increased risk in the subsequent stage should select for switching later and larger. This framework has frequently been applied to metamorphosis in amphibians, with mixed results. Few studies examining the effect of larval predation risk on metamorphosis have observed the predicted pattern, and no studies, to our knowledge, have examined the effect of increased risk during and after metamorphosis on the timing of this switch point. Here we examine the effect of larval and post-metamorphic predation risk on metamorphosis in the red-eyed treefrog, Agalychnis callidryas. We raised tadpoles in the presence or absence of cues from caged water bugs fed larvae and cues from spiders fed emerging metamorphs. Water bugs are effective larval predators, while spiders are poor larval predators but prey on metamorphs. Furthermore, since spiders forage on the water surface it is possible that tadpoles could assess future risk from this predator. Predators induced opposite shifts in life history. Tadpoles emerged smaller and less developed in response to water bugs, but later and larger in response to spiders. Interestingly, predator effects on larval duration were not independent; tadpoles delayed emerging in response to spiders, but only in the absence of water bugs.     

The growth-predation risk trade-off under a growing gape-limited predation threat

Growth is a critical ecological trait because it can determine population demography, evolution, and community interactions. Predation risk frequently induces decreased foraging and slow growth in prey. However, such strategies may not always be favored when prey can outgrow a predator's hunting ability. At the same time, a growing gape-limited predator broadens its hunting ability through time by expanding its gape and thereby creates a moving size refuge for susceptible prey. Here, I explore the ramifications of growing gape-limited predators for adaptive prey growth. A discrete demographic model for optimal foraging/growth strategies was derived under the realistic scenario of gape-limited and gape-unconstrained predation threats. Analytic and numerical results demonstrate a novel fitness minimum just above the growth rate of the gape-limited predator. This local fitness minimum separates a slow growth strategy that forages infrequently and accumulates low but constant predation risk from a fast growth strategy that forages frequently and experiences a high early predation risk in return for lower future predation risk and enhanced fecundity. Slow strategies generally were advantageous in communities dominated by gape-unconstrained predators whereas fast strategies were advantageous in gape-limited predator communities. Results were sensitive to the assumed relationships between prey size and fecundity and between prey growth and predation risk. Predator growth increased the parameter space favoring fast prey strategies. The model makes the testable predictions that prey should not grow at the same rate as their gape-limited predator and generally should grow faster than the fastest growing gape-limited predator. By focusing on predator constraints on prey capture, these results integrate the ecological and evolutionary implications of prey growth in diverse predator communities and offer an explanation for empirical growth patterns previously viewed to be anomalies.     

Adaptive changes in size and age at metamorphosis can qualitatively vary with predator type and available defenses

In many taxa the timing of metamorphosis is plastic in response to predation risk during the pre-metamorphic stage, and trends in both age and body size at metamorphosis have been the subject of much study. The responses to cues of predators are predominantly to be larger or equal-sized at the same age or older at metamorphosis. These observations are in direct contrast with existing theoretical treatments of this plasticity, which mostly predict earlier and smaller metamorphosis and never later and larger metamorphosis without invoking indirect effects on growth rate. Here we resolve the discrepancy between theory and observation using a dynamic state-dependent model that incorporates morphological and behavioral responses to predation risk. We allow prey to choose the optimal activity level and/or investment in defense over the growth period. We show that under certain conditions, metamorphosis at a larger size and later time is likely to be optimal. Our analysis allows us to make testable predictions about the changes in activity level of prey as they grow and how the effect of providing refuges will vary with predator type. Several of these predictions are supported by a meta-analysis of metamorphic responses to caged predators by larval amphibians and insects. Our predictions lead to insights about the feedback effects of antipredator responses on growth and subsequent implications for life history.     

Predator-induced female behavior in the absence of male incubation feeding: an experimental study

Parental care plasticity is critical to understanding the ecological and evolutionary influence of nest predation on life history strategies. In birds, incubation imposes a trade-off between the requirements of females (i.e., food) and egg requirements (i.e., heat and protection from predators). However, studies on this topic are rare and usually restricted to species where the male feeds the incubating female, relaxing her incubation costs. Males and females can reduce their activity at the nest to avoid detection by predators. However, females could follow two alternative antipredator strategies: to delay their return to the nest to avoid attracting attention from the potential predator or to return to the nest as soon as possible to enhance nest concealment. In this study, we manipulated the perceived risk of nest predation of incubating common blackbirds (Turdus merula), a species without incubation feeding, to study female behavioral changes induced by nest predation risk. We show experimentally that female blackbirds can reduce their nest visits in the situation with higher nest predation risk. In addition, we confirm that females significantly delay their return to the nest in the presence of a nest predator, contradicting the nest concealment hypothesis. However, our results could be interpreted as a passive antipredator response (to minimize clues given to predators) or as an active antipredator response (to search for predators to expel them from their territories).     

Sublethal pesticide concentrations and predation jointly shape life history: behavioral and physiological mechanisms.

Despite their relevance for risk assessment, the interactive effects of pesticide and predation cues are poorly understood because the underlying behavioral and physiological mechanisms are largely unknown. To explore these mechanisms, we reared larvae of the damselfly Coenagrion puella at three different predation risk levels and a range of environmentally realistic concentrations of three pesticides used worldwide (atrazine, carbaryl, and endosulfan). We compared key development responses (growth rate, developmental time, and final size) against food ingestion, assimilation, and conversion efficiency, and acetylcholinesterase (AChE) activity. Predation risk impaired all endpoints, including AChE activity, while the effects of pesticide stress were smaller for atrazine and endosulfan and absent for carbaryl. The effects of both stressors and their interaction on life history were mostly indirect through resource acquisition and energy allocation. Compensatory physiological mechanisms to pesticide stress (atrazine and endosulfan) were present in larvae reared in the absence of predation stress but were offset under predation stress. As a result, smaller size (atrazine and endosulfan) and lower growth rate (endosulfan) from pesticide stress were only found in the highest predation risk treatment. Our results provide insight as to the conditions under which interactions between stressors are likely to occur: damselfly populations at high density and living in fish ponds will be more affected by pesticides than populations at low densities in fishless ponds. By identifying variables that may shape the interaction between predation stress and other stressors such as pesticides, our mechanistic approach may help to bridge the gap between laboratory and field studies.     

Evolution of Nesting Height in an Endangered Hawaiian Forest Bird in Response to a Non‐Native Predator

Abstract: The majority of bird extinctions since 1800 have occurred on islands, and non‐native predators have been the greatest threat to the persistence of island birds. Island endemic species often lack life‐history traits and behaviors that reduce the probability of predation and they can become evolutionarily trapped if they are unable to adapt, but few studies have examined the ability of island species to respond to novel predators. The greatest threat to the persistence of the Oahu Elepaio (Chasiempis ibidis), an endangered Hawaiian forest bird, is nest predation by non‐native black rats (Rattus rattus). I examined whether Oahu Elepaio nest placement has changed at the individual and population levels in response to rat predation by measuring nest height and determining whether each nest produced offspring from 1996 to 2011. Average height of Oahu Elepaio nests increased 50% over this 16‐year period, from 7.9 m (SE 1.7) to 12.0 m (SE 1.1). There was no net change in height of sequential nests made by individual birds, which means individual elepaios have not learned to place nests higher. Nests ≤3 m off the ground produced offspring less often, and the proportion of such nests declined over time, which suggests that nest‐building behavior has evolved through natural selection by predation. Nest success increased over time, which may increase the probability of long‐term persistence of the species. Rat control may facilitate the evolution of nesting height by slowing the rate of population decline and providing time for this adaptive response to spread through the population.     

Trade-offs between time, predation risk and life history, and their implications for biogeography: A systems modelling approach with a primate case study

Group sizes are often considered to be the result of a trade-off between predation risk and the costs of feeding competition. We develop a model to explore the interaction between different ecological constraints on group sizes, using a primate (baboons) case study. The model uses climatic correlates of time budgets to predict maximum ecologically tolerable group size, and climatic predictors of predation risk (reflected mainly in predator density and female body mass) to predict minimum tolerable group size for any given habitat. As well as defining the range of sustainable group sizes for a given habitat, the model also allows us to reliably predict our exemplar taxon's biogeographical distribution across Africa. We also explore the life history implications of the model to ask whether baboons form group sizes which maximise survival or fecundity in the classic trade off between these two key life history variables. Our results indicate that, within the range of study sites in our sample, baboons prefer to maximise fecundity. However, the data indicate that in higher predation risk habitats they would switch to maximising survival at the expense of fecundity. We argue that this is due to the fact that interbirth interval and developmental rates have a ceiling that cannot be breached. Thus, while females can shorten interbirth intervals to compensate for increased predation risk, there is a limit to how much these life history variables can be altered, and when this is reached the best strategy is to maximise survivorship.     

Independent effects of food and predator-mediated processes on annual fecundity in a songbird

We investigated the relative importance and interaction of ecological processes affecting annual fecundity in birds by simultaneously manipulating food availability and nest predation risk in a small songbird, the Wrentit (Chamaea fasciata). From 2000 to 2002 we provided supplemental food to individual Wrentit territories, and during 2002 we altered nest predation risk by providing supplemental food to their principal predators, Western Scrub-Jays (Aphelocoma californica). These experiments were conducted during a period of high interannual variation in rainfall, with 2002 being one of the driest years on record. Food-supplemented Wrentits in a normal predation environment produced an average of 0.54 more fledglings per year than control pairs over the three breeding seasons. During the feeding plus predation manipulation experiment, Wrentit food supplementation and lowered nest predation risk each independently increased the probability that a Wrentit pair would fledge young; however, the interaction between food supplementation and altered nest predation risk was not significant. Thus, even in an extreme drought year, both food and nest predation had equal but independent effects on reproductive success and annual fecundity. Combining supplemental food with reduced nest predation did not result in a synergistic increase in annual fecundity, primarily because Wrentits did not produce multiple broods. Our results suggest that whether food and predation have additive or synergistic effects on reproductive success depends on the life history of the species and the environment in which they live.     

Threat-sensitive feeding strategy of immature sticklebacks (Gasterosteus aculeatus) in response to recent experimental infection with the cestode Schistocephalus solidus

Threat-sensitive decision-making might be changed in response to a parasitic infection that impairs future reproduction. Infected animals should take more risk to gain energy to speed up their growth to achieve early reproduction and/or to strengthen their immune response. To avoid correlational evidence, we experimentally infected and sham-infected randomly selected immature three-spined sticklebacks with the cestode Schistocephalus solidus. For 7 weeks we determined the threat-sensitive foraging decisions and growth of individual sticklebacks in the presence of a live pike (Esox lucius). The experimenters were blind with respect to the infection status of the fish. In contrast to previous studies, our recently infected fish should have been almost unconstrained by the parasite and thus have been able to adopt an appropriate life history strategy. We found a strong predator effect for both infected and uninfected fish: the sticklebacks’ risk-sensitive foraging strategy resulted in significantly reduced growth under predation risk. Infected fish did not grow significantly faster under predation risk than uninfected fish. Since infected fish consumed much less prey in the presence of the predator than did infected fish in its absence, they obviously did not use the opportunity to maximize their growth rate to reach reproduction before the parasite impairs it. Received: 21 June 1999 / Revised: 27 November 1999 / Accepted: 5 September 2000     

Toward a predictive theory of risk effects: hypotheses for prey attributes and compensatory mortality

Risk effects, or the costs of antipredator behavior, can comprise a large proportion of the total effect of predators on their prey. While empirical studies are accumulating to demonstrate the importance of risk effects, there is no general theory that predicts the relative importance of risk effects and direct predation. Working toward this general theory, it has been shown that functional traits of predators (e.g., hunting modes) help to predict the importance of risk effects for ecosystem function. Here, I note that attributes of the predator, the prey, and the environment are all important in determining the strength of antipredator responses, and I develop hypotheses for the ways that prey functional traits might influence the magnitude of risk effects. In particular, I consider the following attributes of prey: group size and dilution of direct predation risk, the degree of foraging specialization, body mass, and the degree to which direct predation is additive vs. compensatory. Strong tests of these hypotheses will require continued development of methods to identify and quantify the fitness costs of antipredator responses in wild populations.     

Ecological and hormonal correlates of antipredator behavior in adult Belding’s ground squirrels (Spermophilus beldingi)

Predator–prey relationships provide an excellent opportunity to study coevolved adaptations. Decades of theoretical and empirical research have illuminated the various behavioral adaptations exhibited by prey animals to avoid detection and capture, and recent work has begun to characterize physiological adaptations, such as immune reactions, metabolic changes, and hormonal responses to predators or their cues. A 2-year study quantified the activity budgets and antipredator responses of adult Belding’s ground squirrels (Spermophilus beldingi) living in three different California habitats and likely experiencing different predation pressures. At one of these sites, which is visually closed and predators and escape burrows are difficult to see, animals responding to alarm calls remain alert longer and show more exaggerated responses than adults living in two populations that likely experience less intense predation pressure. They also spend more time alert and less time foraging than adults at the other two sites. A 4-year study using noninvasive fecal sampling of cortisol metabolites revealed that S. beldingi living in the closed site also have lower corticoid levels than adults at the other two sites. The lower corticoids likely reflect that predation risk at this closed site is predictable, and might allow animals to mount large acute cortisol responses, facilitating escape from predators and enhanced vigilance while also promoting glucose storage for the approaching hibernation. Collectively, these data demonstrate that local environments and perceived predation risk influence not only foraging, vigilance, and antipredator behaviors, but adrenal functioning as well, which may be especially important for obligate hibernators that face competing demands on glucose storage and mobilization.     

Uniform predation risk in nature: common,inconspicuous, and a source of error to predation risk experiments

Previous studies indicate that when predation risk is uniform across habitats, foragers concentrate their exploitation in fewer patches. Although uniform predation risk may seem rare in nature, some scenarios might cause it. Testing all scenarios in a single experiment is unfeasible; therefore, we developed a model that points whether concentration of exploitation in specific habitats due to uniform risk requires parameter values similar to what is found in literature. This model was based on Brown’s (Behav Ecol Sociobiol 22:37–47, 1988) fitness function but rescaled to multiple habitats and predators, including uniform risk predators. Deriving function’s maximum allowed comparisons with giving-up density studies. Results showed that uniform predation risk had a u-shaped effect in habitat exploitation, causing a concentration of habitat exploitation at probabilities of survival from 0.2 to 0.8. However, the length of this interval and degree of concentration depended on the value of safety to forager fitness. Heterogeneous, nonuniform, predation risk decreases habitat exploitation where it was higher, therefore suppressing the effect of uniform risk on prey behavior. Time spent in the focal habitat and metabolic costs reduced the detectability of habitat concentration, while total time did not. We also found that uniform risk reduced accuracy of heterogeneous risk measurements. Future studies should aim to control all possible predators, as even the mild ones can induce complex behavior.     

Predator hunting mode and habitat domain alter nonconsumptive effects in predator-prey interactions

Predators can affect prey populations through changes in traits that reduce predation risk. These trait changes (nonconsumptive effects, NCEs) can be energetically costly and cause reduced prey activity, growth, fecundity, and survival. The strength of nonconsumptive effects may vary with two functional characteristics of predators: hunting mode (actively hunting, sit-and-pursue, sit-and-wait) and habitat domain (the ability to pursue prey via relocation in space; can be narrow or broad). Specifically, cues from fairly stationary sit-and-wait and sit-and-pursue predators should be more indicative of imminent predation risk, and thereby evoke stronger NCEs, compared to cues from widely ranging actively hunting predators. Using a meta-analysis of 193 published papers, we found that cues from sit-and-pursue predators evoked stronger NCEs than cues from actively hunting predators. Predator habitat domain was less indicative of NCE strength, perhaps because habitat domain provides less reliable information regarding imminent risk to prey than does predator hunting mode. Given the importance of NCEs in determining the dynamics of prey communities, our findings suggest that predator characteristics may be used to predict how changing predator communities translate into changes in prey. Such knowledge may prove particularly useful given rates of local predator change due to habitat fragmentation and the introduction of novel predators.     

Effects of embryonic exposure to conspecific chemical cues on hatching and larval traits in the common frog <Emphasis Type="Italic">(Rana temporaria)</Emphasis>

Summary. Recent studies indicate that amphibian eggs are capable of hatching plasticity in response to chemical cues released by predators feeding on conspecific eggs or larvae. However, information is scarce on the relative importance of predator and conspecific cues in such a process. In particular, no attempt has been made to compare the effects of embryonic exposures to chemical cues indicative of a predation risk for eggs and larvae, although both life stages can co-occur in natural habitats. In this context, common frog embryos (Rana temporaria) were raised until hatching in the presence of crushed conspecific extracts from eggs and tadpoles to assess their respective influences on some hatching and larval traits. While a significant delay in hatching time was observed in embryos exposed to chemical cues from tadpole extract, this life-history shift appeared unaffected by embryonic exposure to egg extract. Hatchlings derived from eggs incubated in the presence of both conspecific extracts showed a significantly greater weight than unexposed controls. However, such an effect was no longer apparent 15, 30 and 50 days after hatching, suggesting that embryonic exposure to chemical cues from damaged conspecific eggs and tadpoles has no influence on larval growth. Lastly, morphological measurements performed on hatchlings and older tadpoles (15, 30 and 50 days old) revealed no significant effect of embryonic treatments on the shape of body and tail.     

Invasive competitor and native seed predators contribute to rarity of the narrow endemic Astragalus sinuatus Piper

The conservation of rare plant species hinges on our ability to identify the underlying mechanisms that limit rare plant populations. Theory on rarity suggests that both predispersal seed predation and competition can be important mechanisms influencing abundance and/or distribution of rare plant populations. Yet few studies have tested these interactions, and those that have evaluated each mechanism independently. Astragalus sinuatus Piper (Whited's milkvetch) is a narrow endemic plant species restricted to eight populations within a 10-km2 area in eastern Washington. We used experimental and observational methods to test the effects of native insect predispersal seed predators and an invasive grass (Bromus tectorum L. [cheatgrass]) on seed set and population density of A. sinuatus. We quantified per capita seed production and pod predation rates across four sites and among four years. Seed predation rates were high across four sites (66-82%) and all years (65-82%). Experimental reduction of predispersal seed predators significantly increased per capita seed set of A. sinuatus (164-345%) at two experimental sites. Concurrently, two seed addition experiments demonstrated the effect of seed loss and presence of B. tectorum on seedling recruitment and establishment of A. sinuatus over four growing seasons. In the first seed addition experiment, we found no difference in recruitment and establishment between low (40) and high (120) seed addition levels. In the second addition experiment (one level of addition; 40 seeds), we found that recruitment and survivorship increased 200% in plots where B. tectorum was removed compared to plots where B. tectorum was present. Thus, seed addition had no impact in the presence of B. tectorum; conversely, in the absence of B. tectorum, seed addition was highly effective at increasing population numbers. Results suggest that, in areas where B. tectorum is present, recruitment is site limited, and it is seed limited when B. tectorum is absent. We recommend that managers reduce B. tectorum in an effort to increase population growth of A. sinuatus; in areas where B. tectorum is absent, short-term reduction of insect predators should be considered as a strategy to increase population growth of this rare species.     

Antipredator defenses along a latitudinal gradient in Rana temporaria

Antipredator defenses are expected to decrease toward higher latitudes because predation rates are predicted to decrease with latitude. However, latitudinal variation in predator avoidance and defense mechanisms has seldom been studied. We studied tadpole antipredator defenses in seven Rana temporaria populations collected along a 1500-km latitudinal gradient across Sweden, along which previous studies have found increasing tadpole growth and development rates. In a laboratory common garden experiment, we measured behavioral and morphological defenses by raising tadpoles in the presence and absence of a predator (Aeshna dragonfly larva) in two temperature treatments. We also estimated tadpole survival in the presence of free-ranging predators and compared predator densities between R. temporaria breeding ponds situated at low and high latitudes. Activity and foraging were generally positively correlated with latitude in the common garden experiment. While all populations responded to predator presence by decreasing activity and foraging, high-latitude populations maintained higher activity levels in the presence of the predator. All populations exhibited defensive morphology in body and tail shape. However, whereas tail depth tended to increase with latitude in the presence of predator, it did not change with latitude in the absence of the predator. Predator presence generally increased larval period and decreased growth rate. In the southern populations, predator presence tended to have a negative effect on metamorphic size, whereas in the northern populations predators had little or a positive effect on size. Latitude of origin had a strong effect on survival in the presence of a free-ranging predator, with high-latitude tadpoles experiencing higher mortality than those from the low latitudes. In the wild, predator densities were significantly lower in high-latitude than in mid-latitude breeding ponds. Although the higher activity level in the northern populations seems to confer a significant survival disadvantage under predation risk, it is probably needed to maintain the high growth and development rates. However, the occurrence of R. temporaria at high latitudes may be facilitated by the lower predator densities in the north.     

Spatial and temporal changes in group dynamics and range use enable anti-predator responses in African buffalo

The reintroduction of large predators provides a framework to investigate responses by prey species to predators. Considerable research has been directed at the impact that reintroduced wolves (Canis lupus) have on cervids, and to a lesser degree, bovids, in northern temperate regions. Generally, these impacts alter feeding, activity, and ranging behavior, or combinations of these. However, there are few studies on the response of African bovids to reintroduced predators, and thus, there is limited data to compare responses by tropical and temperate ungulates to predator reintroductions. Using the reintroduction of lion (Panthera leo) into the Addo Elephant National Park (AENP) Main Camp Section, South Africa, we show that Cape buffalo (Syncerus caffer) responses differ from northern temperate ungulates. Following lion reintroduction, buffalo herds amalgamated into larger, more defendable units; this corresponded with an increase in the survival of juvenile buffalo. Current habitat preference of buffalo breeding herds is for open habitats, especially during the night and morning, when lion are active. The increase in group size and habitat preference countered initial high levels of predation on juvenile buffalo, resulting in a return in the proportion of juveniles in breeding herds to pre-lion levels. Our results show that buffalo responses to reintroduced large predators in southern Africa differ to those of northern temperate bovids or cervids in the face of wolf predation. We predict that the nature of the prey response to predator reintroduction is likely to reflect the trade-off between the predator selection and hunting strategy of predators against the life history and foraging strategies of each prey species.     

Case selection by a limnephilid caddisfly [Potamophylax latipennis (Curtis)] in response to different predators

Some organisms use morphological structures obtained by behavioural processes to lower mortality by predation. We test whether larvae of the limnephilid caddisfly Potamophylax latipennis (Curtis) vary their responses to the presence of different predators (dragonfly naiads, fire salamander larvae or brown trout) by choosing organic or mineral cases. We offered both case types to larvae, and simulated differences in predation risk using water conditioned with chemicals from the different predators. Our results show that Potamophylax larvae detect and discriminate predators using water-borne chemical cues and alter their choice of case type according to the perceived predation risk. Moreover, the distribution of larvae bearing cases of different anti-predator value matches the spatial variation in predation risk in the field.     

Hunted hunters? Effect of group size on predation risk and growth in the Australian subsocial crab spider Diaea ergandros

A reduced predation risk is considered to be a major adaptive advantage of sociality. While most studies are concerned with non-predatory prey species, group-living predators are likely to face similar threats from higher-order predators. We studied the relationship between group size and predation risk in the subsocial crab spider Diaea ergandros by testing predictions from theoretical models including attack abatement as well as the formation of protective retreats. In a field survey, we found predatory clubionid spiders in 35 % of the D. ergandros nests and as predicted, nest size did not correlate with predator presence. In a subsequent laboratory experiment, we observed survival probability, nest construction activity and feeding behaviour including weight development between groups of different sizes as well as in the absence or presence of a predator. Large groups had an advantage in terms of survival and growth compared to smaller groups or single individuals. They also built significantly larger nests than smaller groups, supporting the idea of protective retreat formation being an adaptive benefit to group living. Even though clubionids did attack D. ergandros, they did not significantly affect overall mortality of D. ergandros. The feeding experiment showed that spiders fed on a larger proportion of flies in the presence of a predator. However, these groups gained significantly less weight compared to the control groups, indicating that the potential predators not only act as predators but also as food competitors, constituting a twofold cost for D. ergandros.