The influence of predator regime on the behaviour and mortality of a freshwater amphipod, <Emphasis Type="Italic">Gammarus pulex</Emphasis>

In species with restricted dispersal, traits may become genetically fixed leading to local adaptations. Therefore, predator avoidance in a prey species may differ between populations experiencing different predator regimes, but also between sexes within a population due to different vulnerability to predators. In this study we used male and female Gammarus pulex from two different predator regimes: fishless ponds, where invertebrates are the dominant predators and ponds with predatory fish. In the laboratory we examined refuge use, mortality, leaf decomposition rate and pair-formation in G. pulex when exposed to predator cues from either invertebrate predators or fish. Individuals from fish ponds spent more time in refuge and had a higher mortality than those from fishless ponds independent of predator cues. There was no effect of pond predator regime or predator cues on leaf decomposition rates. Further, fewer individuals formed pairs in G. pulex from fish ponds than from fishless ponds. Male G. pulex had a higher mortality and a higher decomposition rate than females independent of predator cues. However, there was no difference in refuge use between sexes. Our study shows that there are general differences in behaviour traits, both between predator regimes and sexes in G. pulex.     

Predatory functional response and prey choice identify predation differences between native/invasive and parasitised/unparasitised crayfish

Background

Invasive predators may change the structure of invaded communities through predation and competition with native species. In Europe, the invasive signal crayfish Pacifastacus leniusculus is excluding the native white clawed crayfish Austropotamobius pallipes.

Methodology and Principal Findings

This study compared the predatory functional responses and prey choice of native and invasive crayfish and measured impacts of parasitism on the predatory strength of the native species. Invasive crayfish showed a higher (>10%) prey (Gammarus pulex) intake rate than (size matched) natives, reflecting a shorter (16%) prey handling time. The native crayfish also showed greater selection for crustacean prey over molluscs and bloodworm, whereas the invasive species was a more generalist predator. A. pallipes parasitised by the microsporidian parasite Thelohania contejeani showed a 30% reduction in prey intake. We suggest that this results from parasite-induced muscle damage, and this is supported by a reduced (38%) attack rate and increased (30%) prey handling time.

Conclusions and Significance

Our results indicate that the per capita (i.e., functional response) difference between the species may contribute to success of the invader and extinction of the native species, as well as decreased biodiversity and biomass in invaded rivers. In addition, the reduced predatory strength of parasitized natives may impair their competitive abilities, facilitating exclusion by the invader.     

Opportunistic omnivory impairs our ability to predict invasive species impacts from functional response comparisons

Comparing the relationship between resource use and resource availability (i.e. the functional response, FR) between two predators can provide useful insights on their relative predatory impacts. For instance in invasion ecology, an increase in the predation pressure on local prey populations can be predicted from a significant difference in FR revealing a higher FR for the invasive predator compared to the native trophic analogue it may replace. In traditional FR experiments, the focal prey species is the only source of food. This may lead to misinterpretations with opportunistic omnivores that are able to cope with different resource availabilities in their natural environment, and whose predation rate may therefore be modulated by the presence of alternative resources. To address this question, we compared the FR of two freshwater gammarid species known to behave as opportunistic omnivores: the invasive “killer shrimp” Dikerogammarus villosus and the native Gammarus pulex, in a treatment with a focal prey species as the only food source (the water flea Daphnia magna) and in a treatment with the focal prey and an alternative food source (Carpinus betulus leaves). D. villosus showed a significantly higher FR than G. pulex with water fleas only and providing leaf litter suppressed this difference. The predatory impact of D. villosus might therefore be modulated by the relative availability of live prey compared to the alternative food sources. Increasing the realism of FR experiments through the inclusion of abundant and easily accessible alternative resources, like leaf litter for benthic invertebrates, should refine the predictions made from FR comparisons.     

Differential effects of native vs. invasive predators on a common Caribbean reef fish

Predators may have consumptive (lethal) and non-consumptive (sub-lethal) effects on prey. Non-consumptive effects include altered behavior and reduced growth and fecundity. Native prey may not recognize non-native predators as a threat, and therefore may suffer pronounced effects. Additionally, non-native predators may elicit different behavioral responses from prey compared to native predators. Theory predicts that consumptive effects should be greater for non-native predators (due to prey naiveté), and non-consumptive effects should be greater for native predators (due to predator recognition). To test these hypotheses, I monitored bicolor damselfish (Stegastes partitus) in the presence of invasive predatory Pacific lionfish (Pterois spp.), a native predator (graysby, Cephalopholis cruentata), and an egg predator (bluehead wrasse, Thalassoma bifasciatum). Body size and location of lionfish and graysby were monitored on reefs in the Bahamas. Bicolor fecundity was measured as the number and size of egg-masses that individual fish laid. Bicolor fecundity was negatively correlated with lionfish density but not graysby or bluehead density. Neither predator had a detectable effect on bicolor body size, but lionfish density was negatively correlated with the size of mature adult damselfish. I observed behavioral responses of bicolors to the two piscivores, to bluehead wrasse, and to two herbivorous fishes (Acanthurus coeruleus, Scarus spp.) as non-aggressive controls. Bicolors changed behavior (feeding and aggression) in the presence of all native fishes, but not in the presence of lionfish. Thus, differential effects exist between native and non-native predators, and invasive lionfish pose a non-consumptive threat to bicolor damselfish via reduced growth and fecundity.     

Your worst enemy could be your best friend: predator contributions to invasion resistance and persistence of natives

Native predators are postulated to have an important role in biotic resistance of communities to invasion and community resilience. Effects of predators can be complex, and mechanisms by which predators affect invasion success and impact are understood for only a few well-studied communities. We tested experimentally whether a native predator limits an invasive species’ success and impact on a native competitor for a community of aquatic insect larvae in water-filled containers. The native mosquito Aedes triseriatus alone had no significant effect on abundance of the invasive mosquito Aedes albopictus. The native predatory midge Corethrella appendiculata, at low or high density, significantly reduced A. albopictus abundance. This effect was not caused by trait-mediated oviposition avoidance of containers with predators, but instead was a density-mediated effect caused by predator-induced mortality. The presence of this predator significantly reduced survivorship of the native species, but high predator density also significantly increased development rate of the native species when the invader was present, consistent with predator-mediated release from interspecific competition with the invader. Thus, a native predator can indirectly benefit its native prey when a superior competitor invades. This shows the importance of native predators as a component of biodiversity for both biotic resistance to invasion and resilience of a community perturbed by successful invasion.     

Predicting the ecological impacts of a new freshwater invader: functional responses and prey selectivity of the ‘killer shrimp’, Dikerogammarus villosus,compared to the native Gammarus pulex

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Inter‐specific differences in invader and native fish functional responses illustrate neutral effects on prey but superior invader competitive ability

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Biotic resistance of impact: a native predator (<Emphasis Type="Italic">Chaoborus</Emphasis>) influences the impact of an invasive predator (<Emphasis Type="Italic">Bythotrephes</Emphasis>) in temperate lakes

Introduced predators have caused some of the largest documented impacts of non-native species. Interactions among predators can have complex effects, leading to both synergistic and antagonistic outcomes. Complex interactions with native predators could play an important role in mediating the impact of non-native predators. We explore the role of the native predator context on the effect of the introduced predatory cladoceran Bythotrephes longimanus. While post-invasion impacts have been well described, studies have largely ignored the role of native predators. We used a field mesocosm experiment to determine whether Bythotrephes’ impact on prey communities is influenced by the presence of the ubiquitous native predatory insect larvae Chaoborus. The two predators exhibited niche complementarity as no change in total zooplankton prey abundance was detected across predator treatments. Rather, copepod abundances increased with decreasing abundances of Chaoborus, while cladocerans decreased with increasing abundances of Bythotrephes. Thus, the replacement of Chaoborus with Bythotrephes led to changes in the overall community structure of the zooplankton prey, but had little effect on prey total abundance. More interestingly, we found evidence of biotic resistance of impact, that is, the impact of Bythotrephes on the cladoceran community was altered when the two predators co-occurred. Specifically, the predation effect of Bythotrephes was more restricted to the shallower regions of the water column in the presence of Chaoborus, leading to a reduced impact on deeper dwelling prey taxa. Overall, our results demonstrate that the native predator context is important when trying to understand the effect of non-native predators and that variation in native predator abundances and assemblages could explain variation in impact across invaded habitats.     

Reciprocity in predator–prey interactions: exposure to defended prey and predation risk affects intermediate predator life history and morphology

A vast body of literature exists documenting the morphological, behavioural and life history changes that predators induce in prey. However, little attention has been paid to how these induced changes feed back and affect the predators’ life history and morphology. Larvae of the phantom midge Chaoborus flavicans are intermediate predators in a food web with Daphnia pulex as the basal resource and planktivorous fish as the top predator. C. flavicans prey on D. pulex and are themselves prey for fish; as D. pulex induce morphological defences in the presence of C. flavicans this is an ideal system in which to evaluate the effects of defended prey and top predators on an intermediate consumer. We assessed the impact on C. flavicans life history and morphology of foraging on defended prey while also being exposed to the non-lethal presence of a top fish predator. We tested the basic hypothesis that the effects of defended prey will depend on the presence or absence of top predator predation risk. Feeding rate was significantly reduced and time to pupation was significantly increased by defended morph prey. Gut size, development time, fecundity, egg size and reproductive effort respond to fish chemical cues directly or significantly alter the relationship between a trait and body size. We found no significant interactions between prey morph and the non-lethal presence of a top predator, suggesting that the effects of these two biological factors were additive or singularly independent. Overall it appears that C. flavicans is able to substantially modify several aspects of its biology, and while some changes appear mere consequences of resource limitation others appear facultative in nature.     

Pre-adaptive shift of a native predator (Araneae, Zodariidae) to an abundant invasive ant species (Hymenoptera, Formicidae)

Invasive species often displace native species and can affect ecological processes in invaded habitats. If invasive species become abundant, changes in prey availability may be particularly harmful to specialist predators. The Argentine ant, Linepithema humile Mayr, is an important invasive species on nearly all continents. Spiders of the genus Zodarion are specialised ant-eating predators native to the Mediterranean yet it is unknown if they can exploit invasive ant species. Here we studied spatial and temporal abundance of this invasive ant and the native spider, Zodarion cesari Pekár, during 4?years in four citrus groves. Circadian activity of both spiders and ants, and capture efficiency and prey specificity of the predator were also evaluated. The abundance of Z. cesari was strongly correlated to L. humile abundance. The predatory activity of spiders varied seasonally with differences on the relative frequency of spiders capturing ants depending on the time of the year. In laboratory, Z. cesari displayed most efficient capture upon the native ant Tapinoma nigerrimum (Nylander) and the invasive ant L. humile in comparison with five other native ant species. These results demonstrate that the native spider Z. cesari is successfully exploiting the invasive ant species L. humile and is likely a locally monophagous predator. We suggest that Z. cesari shifted away from native T. nigerrimum post invasion as both ant species are phylogenetically related and of similar size.     

Modeling the impacts of global warming on predation and biotic resistance: mosquitoes, damselflies and avian malaria in Hawaii

Biotic resistance from native predators can play an important role in regulating or limiting exotic prey. We investigate how global warming potentially alters the strength and spatial extent of these predator–prey interactions in aquatic insect ecosystems. As a simple model system, we use rock pools in streams of rainforests of Hawaii, which contain the beautiful Hawaiian damselfly Megalagrion calliphya as predator and the invasive southern house mosquito Culex quinquefasciatus as prey. This abundant mosquito is the major vector of avian malaria transmission to native forest birds. We use mathematical modeling to evaluate the potential impacts of damselfly predation and temperature on mosquito population dynamics. We model this predator–prey system along an elevational gradient (749-1952 m elevation) and assess the effect of 1°C and 2°C climate warming scenarios as well as the effects of El Niño and La Niña oscillations, on predator–prey dynamics. Our results indicate that the strength of biotic resistance of native predators on invasive prey may decrease with increasing temperature because demographic rates of predator and prey are differentially affected by temperature. Future warming could therefore increase the abundance of invasive species by releasing them from predation pressure. If the invasive species is a disease vector, these shifts could increase the impact of disease on both humans and wildlife.     

The functional role of Gammarus(Crustacea, Amphipoda): shredders, predators, or both?

Gammarus spp. are traditionally viewed under the functional feeding group (FFG) concept as herbivorous `shredders'. Although recent studies suggest that Gammarus should also be viewed as predators, this latter role remains contentious. Here, in a laboratory experiment, we objectively examine the balance between shredder and predator roles in a common freshwater species. Gammarus pulex preyed significantly on mayfly nymph, Baetis rhodani, in both the presence and absence of excess leaf material. There was no significant difference in predation where the alternative food, that is, leaf material, was present as compared to absent. Also, G. pulex shredded leaf material in the presence and absence of B. rhodani. However, shredding was significantly reduced where alternative food, that is, B. rhodani prey, was present as compared to absent. Further, G. pulex had a clear leaf species preference. Our results suggest that Gammarus function as both predators and shredders, with the balance of the two roles perhaps depending on food availability and quality. We discuss implications for the use of the FFG concept in assessing freshwater processes, and the role that Gammarus predation may play in structuring macroinvertebrate communities.     

Predator–prey interactions and metabolic rates are altered in stable and unstable groups in a social fish

Understanding the determinants and consequences of predation effort, success and prey responses is important since these factors affect the fitness of predators and prey. When predators are also invasive species, the impacts on prey can be particularly far-reaching with ultimate ecosystem-level consequences. However, predators are typically viewed as behaviourally fixed within this interaction and it is unclear how variation in predator social dynamics affects predator–prey interactions. Using the invasive eastern mosquitofish Gambusia holbrooki and a native glass shrimp Paratya australiensis in Australia, we investigated how varying levels of social conflict within predator groups influences predator–prey interactions. By experimentally manipulating group stability of G. holbrooki, we show that rates of social conflict were lower in groups with large size differences, but that routine metabolic rates were higher in groups with large size differences. Predation effort and success did not vary depending on group stability, but in stable groups predation effort by aggressive dominants was greater than subordinates. The anti-predator responses of prey to the stability of predator groups were mixed. While more prey utilized shelters when exposed to stable compared to unstable groups of predators, a greater proportion were sedentary when predator groups were unstable. Overall, this study demonstrates predator group stability is modulated by differences in body size and can influence prey responses. Further, it reveals a hidden metabolic cost of living in stable groups despite reduced overt social conflict. For invasive species management, it is therefore important to consider the behavioural and physiological plasticity of the invasive predators, whose complex social interactions and metabolic demands can modulate patterns of predator–prey interactions.     

Predator impacts on stream benthic prey

The impact that predators have on benthic, macroinvertebrate prey density in streams is unclear. While some studies show a strong effect of predators on prey density, others show little or no effect. Two factors appear to influence the detection of predator impact on prey density in streams. First, many field studies have small sample sizes and thus might be unable to detect treatment effects. Second, streams contain two broad classes of predators, invertebrates and vertebrates, which might have different impacts on prey density for a variety of reasons, including availability of refuge for prey and prey emigration responses to the two types of predators. In addition, predatory vertebrates have more complex prey communities than predatory invertebrates; this complexity might reduce the impact that predatory vertebrates have on prey because of indirect effects. I conducted a meta-analysis on the results of field studies that manipulate predator density in enclosures to determine (1) if predators have a significant impact on benthic prey density in streams, (2) if the impacts that predatory invertebrates and vertebrates have differ, and (3) if predatory vertebrates have different impacts on predatory prey versus herbivorous prey. The results of the meta-analysis suggest that on average predators have a significant negative effect on prey density, predatory invertebrates have a significantly stronger impact than predatory vertebrates, and predatory vertebrates do not differ in their impact on predatory versus herbivorous invertebrate prey. Three methodological variables (mesh size of enclosures, size of enclosures, and experimental duration) were examined to determine if cross correlations exist that may explain the differences in impact between predatory invertebrates and vertebrates. No correlation exists between mesh size and predator impact. Over all predators, no correlation exists between experimental duration and predator impact; however, within predatory invertebrates a correlation does exist between these variables. Also, a correlation was found between enclosure size and predator impact. This correlation potentially explains the difference in impact between predatory invertebrates and predatory vertebrates. Results of the meta-analysis suggest two important areas for future research: (1) manipulate both types of predators within the same system, and (2) examine their impacts on the same spatial scale.     

The influence of prey identity and size on selection of prey by two marine fishes

Laboratory experiments were conducted with two predatory fishes, Lagodon rhomboides (Linnaeus) and Syngnathus floridae (Jordan & Gilbert), to determine if prey selection was a result of predator preference or prey accessibility. Prey consisted of two species of caridean shrimp, Tozeuma carolinense (Kingsley) and Hippolyte zostericola (Smith), that commonly inhabit seagrass meadows. Natural diets of both predators revealed that selection ofTozeuma and Hippolyte was not related to their field densities. My experiments demonstrated that natural prey selection was a consequence of prey accessibility, not predator preference. Experiments examining the role of prey size in predator preference revealed that large individuals were significantly preferred over small individuals. Observations of prey behavior indicated that prey motion affected predator choice. These results suggest that predator preference was primarily determined by prey visibility and that the combined effects of prey size and motion contributed to the visibility of these prey.     

Reduced drift activity of two benthic invertebrate species is mediated by infochemicals of benthic fish

Regulating mobility by actively entering the drift under imminent predation risk is an avoidance strategy employed by aquatic macroinvertebrate species that is widely accepted within the scientific community. This response was most evident with respect to diurnal predators that feed in the water column, such as many salmonids. We investigated the role of the nocturnal benthivorous gudgeon [Gobio gobio (L.)] on the drift activity of two macroinvertebrate species known to display this behaviour: Baetis rhodani (PICTET) and Gammarus pulex (L.). Laboratory drift experiments using gudgeon kairomones were conducted with the results determining significant altered activity in the presence of gudgeon kairomones for both macroinvertebrate species. B. rhodani showed reduced drift activity in the kairomone treatment compared to the kairomone-free control, with a distinct nocturnal pattern being observed for both. G. pulex shifted from a similar day/night movement pattern to a nocturnal movement pattern with decreased activity during the day. Reduce activity during the day, whilst maintaining normal activity at night would not reduce the probability of encountering a nocturnal predator under natural conditions and therefore appears to not be a meaningful anti-predator response. To assess the relevance of these findings under natural conditions, we compared the experimental results with drift measurements from field observations. These show a significant reduction in drift activity for G. pulex and slight tendencies for reduced night-time drift for B. rhodani, under seasonal variations. We conclude that the behaviour in response to the physical contact or the hydrodynamic stimuli of nocturnal predators is the most likely explanation for the differences between the results from our laboratory experiment and the field observation. We further discuss that the observed migration patterns might have different species specific consequences for density stabilisation on a population level.     

Independent and combined effects of multiple predators across ontogeny of a dominant grazer

Ecosystems host multiple coexisting predator species whose interactions may strengthen or weaken top–down control of grazers. Grazer populations often exhibit size‐structure, but the nature of multiple predator effects on suppression of size‐structured prey has seldom been explicitly considered. In a southeastern US salt‐marsh, we used both field (additive design) and mesocosm (additive‐substitutive design) experiments to test the independent and combined effects of two species of predatory crab on the survival and predator‐avoidance behavior (i.e. a non‐consumptive effect) of both juveniles and adults of a dominant grazing snail. Results showed: 1) juvenile snails were more vulnerable to predation; 2) consumptive impacts of predators were hierarchically nested, i.e. the larger predator consumed both juvenile and adult snails, while the smaller‐bodied predator consumed only juvenile snails; 3) there were no emergent multiple predator effects on snail consumption; and 4) non‐consumptive effects differed from consumptive effects, with only the large predator inducing predator‐avoidance behavior of individuals within either snail ontogenetic class. The smaller predator therefore played a functionally redundant trophic role across the prey classes considered, augmenting and potentially stabilizing trophic regulation of juvenile snails. Meanwhile, the larger predator played a complementary and functionally unique role by both expanding the size‐spectrum of prey trophic regulation and non‐consumptively altering prey behavior. While our study suggests that nestedness of consumptive interactions determined by predator and prey body sizes may allow prediction of the functional redundancy of particular predator species, it also shows that traits beyond predator body size (e.g. habitat domain) may be required to predict potentially cascading non‐consumptive effects. Future studies of multiple predators (and predator biodiversity) should continue to strive towards greater realism by incorporating not only size‐structured prey, but also other aspects of resource and environmental heterogeneity typical of natural ecosystems.     

Feedback between size balance and consumption strongly affects the consequences of hatching phenology in size‐dependent predator–prey interactions

In many size‐dependent predator–prey systems, hatching phenology strongly affects predator–prey interaction outcomes. Early‐hatched predators can easily consume prey when they first interact because they encounter smaller prey. However, this process by itself may be insufficient to explain all predator–prey interaction outcomes over the whole interaction period because the predator–prey size balance changes dynamically throughout their ontogeny. We hypothesized that hatching phenology influences predator–prey interactions via a feedback mechanism between the predator–prey size balance and prey consumption by predators. We experimentally tested this hypothesis in an amphibian predator–prey model system. Frog tadpoles Rana pirica were exposed to a predatory salamander larva Hynobius retardatus that had hatched 5, 12, 19 or 26 days after the frog tadpoles hatched. We investigated how the salamander hatch timing affected the dynamics of prey mortality, size changes of both predator and prey, and their subsequent life history (larval period and size at metamorphosis). The predator–prey size balance favoured earlier hatched salamanders, which just after hatching could successfully consume more frog tadpoles than later hatched salamanders. The early‐hatched salamanders grew rapidly and their accelerated growth enabled them to maintain the predator‐superior size balance; thus, they continued to exert strong predation pressure on the frog tadpoles in the subsequent period. Furthermore, frog tadpoles exposed to the early‐hatched salamanders were larger at metamorphosis and had a longer larval period than other frog tadpoles. These results suggest that feedback between the predator‐superior size balance and prey consumption is a critical mechanism that strongly affects the impacts of early hatching of predators in the short‐term population dynamics and life history of the prey. Because consumption of large nutrient‐rich prey items supports the growth of predators, a similar feedback mechanism may be common and have strong impacts on phenological shifts in size‐dependent trophic relationships.     

Recognition and Response to Native and Novel Predators in the Largespring mosquitofish,Gambusia geiseri

The introduction of predator species into new habitats is an increasingly common consequence of human activities, and the persistence of native prey species depends upon their response to these novel predators. In this study, we examined whether the Largespring mosquitofish, Gambusia geiseri exhibited antipredator behavior and/or an elevation of circulating stress hormones (cortisol) to visual and chemical cues from a native predator, a novel predator, or a non‐predatory control fish. Prey showed the most pronounced antipredator response to the native predator treatment, by moving away from the stimulus, while the prey showed no significant changes in their vertical or horizontal position in response to the novel or non‐predator treatments. We also found no significant difference in water‐borne cortisol release rates following any of the treatments. Our results suggest the prey did not recognize and exhibit antipredator behavior to the novel predator, and we infer that this predator species could be detrimental if it expands into the range of this prey species. Further, our study demonstrates prey may not respond to an invasive predator that is phylogenetically, behaviorally, and morphologically dissimilar from the prey species' native predators.     

Size matters: predation of fish eggs and larvae by native and invasive amphipods

Invasive predators can have dramatic impacts on invaded communities. Extreme declines in macroinvertebrate populations often follow killer shrimp (Dikerogammarus villosus) invasions. There are concerns over similar impacts on fish through predation of eggs and larvae, but these remain poorly quantified. We compare the predatory impact of invasive and native amphipods (D. villosus and Gammarus pulex) on fish eggs and larvae (ghost carp Cyprinus carpio and brown trout Salmo trutta) in the laboratory. We use size-matched amphipods, as well as larger D. villosus reflecting natural sizes. We quantify functional responses, and electivity amongst eggs or larvae and alternative food items (invertebrate, plant and decaying leaf). D. villosus, especially large individuals, were more likely than G. pulex to kill trout larvae. However, the magnitude of predation was low (seldom more than one larva killed over 48 h). Trout eggs were very rarely killed. In contrast, carp eggs and larvae were readily killed and consumed by all amphipod groups. Large D. villosus had maximum feeding rates 1.6–2.0 times higher than the smaller amphipods, whose functional responses did not differ. In electivity experiments with carp eggs, large D. villosus consumed the most eggs and the most food in total. However, in experiments with larvae, consumption did not differ between amphipod groups. Overall, our data suggest D. villosus will have a greater predatory impact on fish populations than G. pulex, primarily due to its larger size. Higher invader abundance could amplify this difference. The additional predatory pressure could reduce recruitment into fish populations.