PHENOTYPIC DIVERSIFICATION ACROSS AN ENVIRONMENTAL GRADIENT: A ROLE FOR PREDATORS AND RESOURCE AVAILABILITY ON THE EVOLUTION OF LIFE HISTORIES

Changes in age/size‐specific mortality, due to such factors as predation, have potent evolutionary consequences. However, interactions with predators commonly impact prey growth rates and food availability and such indirect effects may also influence evolutionary change. We evaluated life‐history differences in Trinidadian killifish, Rivulus hartii, across a gradient in predation. Rivulus are located in (1) “high predation” sites with large piscivores, (2) “Rivulus/guppy” sites with guppies, and (3) “Rivulus‐only” sites with just Rivulus. Rivulus suffer higher mortality with large predators, and guppies may prey upon small/young Rivulus in Rivulus/guppy environments. In turn, population densities decline while growth rates increase in both localities compared to Rivulus‐only sites. To explore how the direct and indirect effects of predators and guppies influence trait diversification in Rivulus, we examined life‐history phenotypes across five rivers. High predation phenotypes exhibited a smaller size at reproduction, a greater number of eggs that were smaller, and increased reproductive allotment. Such changes are consistent with a direct response to predation. Rivulus from Rivulus/guppy sites were intermediate; they exhibited a smaller size at reproduction, increased fecundity, smaller eggs, and larger reproductive allotment than Rivulus‐only fish. These changes are consistent with models that incorporate the impacts of growth and resources.     

Increased juvenile predation is not associated with evolved differences in adult brain size in Trinidadian killifish (Rivulus hartii)

Vertebrates exhibit extensive variation in brain size. The long‐standing assumption is that this variation is driven by ecologically mediated selection. Recent work has shown that an increase in predator‐induced mortality is associated with evolved increases and decreases in brain size. Thus, the manner in which predators induce shifts in brain size remains unclear. Increased predation early in life is a key driver of many adult traits, including life‐history and behavioral traits. Such results foreshadow a connection between age‐specific mortality and selection on adult brain size. Trinidadian killifish, Rivulus hartii, are found in sites with and without guppies, Poecilia reticulata. The densities of Rivulus drop dramatically in sites with guppies because guppies prey upon juvenile Rivulus. Previous work has shown that guppy predation is associated with the evolution of adult life‐history traits in Rivulus. In this study, we compared second‐generation laboratory‐born Rivulus from sites with and without guppies for differences in brain size and associated trade‐offs between brain size and other components of fitness. Despite the large amount of existing research on the importance of early‐life events on the evolution of adult traits, and the role of predation on both behavior and brain size, we did not find an association between the presence of guppies and evolutionary shifts in Rivulus brain size. Such results argue that increased rates of juvenile mortality may not alter selection on adult brain size.     

LIFE-HISTORY EVOLUTION IN GUPPIES (POECILIA RETICULATA): 1. PHENOTYPIC AND GENETIC CHANGES IN AN INTRODUCTION EXPERIMENT

Previous investigations (Reznick and Endler, 1982; Reznick, 1982a, 1982b) demonstrated that genetic differences in guppy life histories were associated with differences in predation. Guppies from localities with the pike cichlid Crenicichla alta and associated predators matured earlier, had greater reproductive efforts, and produced more and smaller offspring than did guppies from localities with only Rivulus harti as a potential predator. Crenicichla preys primarily on large, sexually mature size-classes of guppies, while Rivulus preys primarily on small, immature size-classes. These patterns of predation are hypothesized to alter mean age-specific survival. Theoretical treatments of such differences in survival predict the observed trends in age at maturity and reproductive effort. We are using introduction experiments to evaluate the role of predators in selecting for these life-history patterns. The experiment whose results are presented here was conducted in a tributary to the El Cedro River (Trinidad), where a waterfall was the upstream limit to the distribution of all fish except Rivulus. Guppies collected from the Crenicichla locality immediately below the waterfall (the downstream control) were introduced over the waterfall in 1981. This introduction released the guppies from Crenicichla predation, exposed them instead to Rivulus predation only, and also introduced them to a different environment, since the introduction site has greater canopy cover than the site of origin. Changes in guppy life-history patterns can be attributed to predation and/or the environment. Evidence from fish collected and preserved in the field demonstrated that, by mid-1983, guppies from the introduction site above the waterfall matured at larger sizes and produced fewer, larger offspring. There were no consistent differences in reproductive allotment (weight of offspring/total weight). With the exception of reproductive allotment, these patterns are identical to previous comparisons between Rivulus and Crenicichla localities. A laboratory genetics experiment demonstrated that males from the introduction site matured at a later age and at a larger size than did males from the control site downstream, as predicted from the “age-specific predation” hypothesis. No differences between localities were observed for female age and size at maturity or for reproductive effort. The trends for fecundity and offspring size were the reverse of those observed in the field. Because only the males changed in the predicted fashion, it is not possible either to reject or to accept the hypothesis of age-specific predation at this time. We discuss the possible causes for these patterns and the high degree of plasticity in the life history, as evidenced by the differences in fecundity and offspring size between the field and laboratory results.     

LIFE-HISTORY EVOLUTION IN GUPPIES: 2. REPEATABILITY OF HELD OBSERVATIONS AND THE EFFECTS OF SEASON ON LIFE HISTORIES

Natural populations of guppies that co-occur with the pike cichlid Crenicichla alta and associated predators mature at smaller body sizes, produce more and smaller offspring per litter reproduce more frequently, and have higher reproductive allotments (weight of developing embryos/total body weight) than guppies that co-occur with just the killifish Rivulus harti (Reznick and Endler, 1982). I here consider three forms of repeatability in these life-history patterns: i) among replicate samples collected on the same day from the same locality, ii) between Crenicichla and Rivulus communities among a new series of localities, and iii) among a smaller series of Crenicichla and Rivulus localities sampled in two wet and two dry seasons. In the analysis of replicate collections from two localities, seven of eight statistical comparisons revealed no significant difference. The usual methodology for estimating these variables therefore accurately represents guppy life-history patterns at a given locality. Differences among guppies from Rivulus and Crenicichla localities, covering a wider geographical area than considered by Reznick and Endler (1982), were virtually identical to the previous comparison. Wet-season samples were associated with significant decreases in reproductive allotment and fecundity and significant increases in the size of mature males and the minimum size of reproducing females. Differences between guppies from Rivulus and Crenicichla localities persisted across all samples and were consistent with all other observations, although they tended to be smaller during the wet season. Discriminant analyses on female reproductive traits showed that fecundity and offspring size made strong, independent contributions to discriminating between guppies from the two types of localities. The contribution from reproductive allotment was considerably smaller. There was more overlap between predator treatments during the wet season. Only 8.5% of the individuals were misclassified during the dry season, but 19.5% were misclassified during the wet season.     

Experimentally induced life-history evolution in a killifish in response to the introduction of guppies

Life-history theory predicts that increased predation on juvenile age/size-classes favors delayed maturation and decreased reproductive investment. Although this theory has received correlative support, experimental tests in nature are rare. In 1976 and 1981, guppies (Poecilia reticulata) were transplanted into localities that previously only contained a killifish, Rivulus hartii. This situation presents an opportunity to experimentally test this life-history prediction because guppies prey upon young Rivulus. We evaluated the response to selection in Rivulus by measuring phenotypic and genotypic divergence between introduction and upstream "control" localities that lack guppies. Contrary to expectations, Rivulus from the introduction sites evolved earlier maturation and increased reproductive investment within 25 years. Such evolutionary changes parallel previous investigations on natural communities of Rivulus, but do not comply with predictions of age/size-specific theory. Guppies also caused reduced densities and increased growth rates of Rivulus, which are hypothesized indirect effects of predation. Additional life-history theories show that changes in density and growth can interact with predator-induced mortality to alter the predicted trajectory of evolution. We discuss how these latter frameworks improve the fit between theory and evolution in Rivulus.     

The evolution of vertebrate eye size across an environmental gradient: phenotype does not predict genotype in a Trinidadian killifish

Vertebrates exhibit substantial variation in eye size. Eye size correlates positively with visual capacity and behaviors that enhance fitness, such as predator avoidance. This foreshadows a connection between predation and eye size evolution. Yet, the conditions that favor evolutionary shifts in eye size, besides the well‐known role for light availability, are unclear. We tested the influence of predation on the evolution of eye size in Trinidadian killifish, Rivulus hartii. Rivulus are located across a series of communities where they coexist with visually oriented piscivores ("high predation" sites), and no predators (“Rivulus‐only” sites). Wild‐caught Rivulus from high predation sites generally exhibited a smaller relative eye size than communities that lack predators. Yet, such differences were inconsistent across rivers. Second‐generation common garden reared fish revealed repeatable decreases in eye size in Rivulus from high predation sites. We performed additional experiments that tested the importance of light and resources on eye size evolution. Sites that differ in light or resource availability did not differ in eye size. Our results argue that differences in predator‐induced mortality underlie genetically‐based shifts in vertebrate eye size. We discuss the drivers of eye size evolution in light of the nonparallel trends between the phenotypic and common garden results.     

An experimental test of antagonistic effects of competition and parasitism on host performance in semi‐natural mesocosms

The mechanisms by which parasites can mediate the interactions between species have received increased interest in recent years. Nonetheless, most research has focused on the role of shared parasites as mediators of interspecific competition. Here, we explore the relative effects of Gyrodactylus specialist ectoparasites of Trinidadian guppies Poecilia reticulata on competition between their host and juveniles of the killifish Rivulus hartii. In mesocosms that replicate natural streams, we exposed guppies to only competitors, to only parasites, to both parasites and competitors, or the absence of both. Consistent with previous studies, we found that female guppies grew significantly less where only Gyrodactylus were present, and this was regardless of infection status or parasite load. Surprisingly, this effect of Gyrodactylus on the growth of female guppies was greatly reduced when both parasites and competitors were present in the mesocosms. We conclude that guppies can mediate the effects of Gyrodactylus on competition with Rivulus, by adaptively fine‐tuning their phenotype when simultaneously facing multiple enemies.     

The relevance of ants as seed rescuers of a primarily bird-dispersed tree in the Neotropical cerrado savanna

Sex ratios can influence mating behaviour, population dynamics and evolutionary trajectories; yet the causes of natural sex ratio variation are often uncertain. Although secondary (birth) sex ratios in guppies (Poecilia reticulata) are typically 1:1, we recorded female-biased tertiary (adult) sex ratios in about half of our 48 samples and male-biased sex ratios in none of them. This pattern implies that some populations experience male-biased mortality, perhaps owing to variation in predation or resource limitation. We assessed the effects of predation and/or inter-specific resource competition (intraguild predation) by measuring the local catch-per-unit-effort (CPUE) of species (Rivulus killifish and Macrobrachium prawns) that may differentially prey on male guppies. We assessed the effects of resource levels by measuring canopy openness and algal biomass (chlorophyll a concentration). We found that guppy sex ratios were increasingly female-biased with increasing CPUE of Macrobrachium, and perhaps also Rivulus, and with decreasing canopy openness. We also found an interaction between predators and resource levels in that the effect of canopy openness was greatest when Macrobrachium CPUE was highest. Our study thus also reveals the value of simultaneously testing multiple environmental factors that may drive tertiary sex ratio variation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Sensory cues of a top‐predator indirectly control a reef fish mesopredator

Behavioural trophic cascades highlight the importance of indirect/risk effects in the maintenance of healthy trophic‐level links in complex ecosystems. However, there is limited understanding on how the loss of indirect top–down control can cascade through the food‐web to modify lower level predator–prey interactions. Using a reef fish food‐web, our study examines behavioural interactions among predators to assess how fear elicited by top‐predator cues (visual and chemical stimuli) can alter mesopredator behaviour and modify their interaction with resource prey. Under experimental conditions, the presence of any cue (visual, chemical, or both) from the top‐predator (coral trout Plectropomus leopardus) strongly restricted the distance swum, area explored and foraging activity of the mesopredator (dottyback Pseudochromis fuscus), while indirectly triggering a behavioural release of the resource prey (recruits of the damselfish Pomacentrus chrysurus). Interestingly, the presence of a large non‐predator species (thicklip wrasse Hemigymnus melapterus) also mediated the impact of the mesopredator on prey, as it provoked mesopredators to engage in an ‘inspection’ behaviour, while significantly reducing their feeding activity. Our study describes for the first time a three‐level behavioural cascade of coral reef fish and stresses the importance of indirect interactions in marine food‐webs.     

Life-history evolution in guppies. VII. The comparative ecology of high- and low-predation environments

Prior research has demonstrated a strong association between the species of predators that co-occur with guppies and the evolution of guppy life histories. The evolution of these differences in life histories has been attributed to the higher mortality rates experienced by guppies in high-predation environments. Here, we evaluate whether there might be indirect effects of predation on the evolution of life-history patterns and whether there are environmental differences that are correlated with predation. To do so, we quantified features of the physical and chemical environment and the population biology of guppies from seven high- and low-predation localities. We found that high-predation environments tend to be larger streams with higher light levels and higher primary productivity, which should enhance food availability for guppies. We also found that guppy populations from high-predation environments have many more small individuals and fewer large individuals than those from low-predation environments, which is caused by their higher birth rates and death rates. Because of these differences in size distribution, guppies from high-predation environments have only one-fourth of the biomass per unit area, which should also enhance food availability for guppies in these localities. Guppies from high-predation sites allocate more resources to reproduction, grow faster, and attain larger asymptotic sizes, all of which are consistent with higher levels of resource availability. We conclude that guppies from high-predation environments experience higher levels of resource availability in part because of correlated differences in the environment (light levels, primary productivity) and in part as an indirect consequence of predation (death rates and biomass density). These differences in resource availability can, in turn, augment the effect of predator-induced mortality as factors that shape the evolution of guppy life-history patterns. We found no differences in the invertebrate communities from high- and low-predation localities, so we conclude that there do not appear to be multitrophic, indirect effects associated with these differences in predation.     

Predators can influence the host‐parasite dynamics of their prey via nonconsumptive effects

Ecological communities are partly structured by indirect interactions, where one species can indirectly affect another by altering its interactions with a third species. In the absence of direct predation, nonconsumptive effects of predators on prey have important implications for subsequent community interactions. To better understand these interactions, we used a Daphnia‐parasite‐predator cue system to evaluate if predation risk affects Daphnia responses to a parasite. We investigated the effects of predator cues on two aspects of host–parasite interactions (susceptibility to infection and infection intensity), and whether or not these effects differed between sexes. Our results show that changes in response to predator cues caused an increase in the prevalence and intensity of parasite infections in female predator‐exposed Daphnia. Importantly, the magnitude of infection risk depended on how long Daphnia were exposed to the cues. Additionally, heavily infected Daphnia that were constantly exposed to cues produced relatively more offspring. While males were ~5× less likely to become infected compared to females, we were unable to detect effects of predator cues on male Daphnia–parasite interactions. In sum, predators, prey, and their parasites can form complex subnetworks in food webs, necessitating a nuanced understanding of how nonconsumptive effects may mediate these interactions.     

Indirect food web interactions increase growth of an algivorous stream fish

1. We tested the hypothesis that indirect food web interactions between some common, invertivorous fishes and their prey would positively affect growth of an algivorous fish species. Specifically, we predicted that orangethroat darter (Etheostoma spectabile) would increase periphyton biomass via a top‐down pathway, indirectly enhancing growth of the algivorous central stoneroller minnow (Campostoma anomalum). Moreover, we predicted that sand shiner (Notropis stramineus) would increase periphyton biomass via a bottom‐up pathway and indirectly enhance growth of the stoneroller minnow. 2. In an 83‐day experiment in large, outdoor, stream mesocosms, we stocked two fish species per mesocosm (stoneroller and either darter or shiner), estimated the effects of the invertivorous and grazing fishes on periphyton biomass and estimated growth of the algivorous fish. 3. The darter consumed grazing invertebrates, indirectly increasing periphyton biomass. The shiner consumed terrestrial insects as predicted, but it did not affect periphyton biomass. 4. In support of our hypothesis, the darter indirectly enhanced stoneroller growth. As predicted, stonerollers consumed the increased periphyton in streams with darters, resulting in greater growth, condition and gut fullness compared to streams without darters. No indirect interaction was observed between stonerollers and shiners. 5. Our study suggests that some invertivorous fish species can positively affect growth of algivorous fishes through indirect food web interactions. Thus, in stream communities, it is possible that the loss of a single, invertivorous fish taxon could have negative consequences on algivorous fish populations via the removal of positive indirect food web interactions.     

Food availability as a major driver in the evolution of life‐history strategies of sibling species

Life‐history theory predicts trade‐offs between reproductive and survival traits such that different strategies or environmental constraints may yield comparable lifetime reproductive success among conspecifics. Food availability is one of the most important environmental factors shaping developmental processes. It notably affects key life‐history components such as reproduction and survival prospect. We investigated whether food resource availability could also operate as an ultimate driver of life‐history strategy variation between species. During 13 years, we marked and recaptured young and adult sibling mouse‐eared bats (Myotis myotis and Myotis blythii) at sympatric colonial sites. We tested whether distinct, species‐specific trophic niches and food availability patterns may drive interspecific differences in key life‐history components such as age at first reproduction and survival. We took advantage of a quasi‐experimental setting in which prey availability for the two species varies between years (pulse vs. nonpulse resource years), modeling mark‐recapture data for demographic comparisons. Prey availability dictated both adult survival and age at first reproduction. The bat species facing a more abundant and predictable food supply early in the season started its reproductive life earlier and showed a lower adult survival probability than the species subjected to more limited and less predictable food supply, while lifetime reproductive success was comparable in both species. The observed life‐history trade‐off indicates that temporal patterns in food availability can drive evolutionary divergence in life‐history strategies among sympatric sibling species.     

Life history trait divergence among populations of a non‐palatable species reveals strong non‐trophic indirect effects of an abundant herbivore

When large herbivores exert selection on their prey plant species, co‐occurring, non‐prey species may experience selection through non‐trophic indirect effects. Such selection is likely common where herbivores are overabundant. Yet, empirical studies of non‐trophic indirect effects as drivers of non‐prey trait evolution are lacking. Here we test for adaptive shifts in life history traits in an unpalatable species, Arisaema triphyllum, a common forest perennial that is unique because it exhibits size‐dependent sex switching. We collected A. triphyllum from six sites that experience a gradient in abiotic stress caused by deer browse pressure on prey plant species that generate indirect effects. We grew A. triphyllum from these sites in a common garden for five years to evaluate life history predictions linking strong indirect effects and abiotic stress to changes in life history traits: flowering onset size threshold, female flowering size threshold, relative growth rate (RGR), biomass allocation, and asexual reproduction. Despite observed differences among phenotypes in the field, expression of flowering onset size threshold, biomass allocation, and asexual reproduction did not differ among the six populations in the garden, indicating common plastic responses. In contrast, A. triphyllum collected from sites experiencing the two highest deer impacts exhibited smaller female flowering size thresholds and the highest RGR. Responses in these traits support the predictions of adaptive divergence in response to indirect effects. Our results reinforce the idea that non‐trophic indirect effects of large herbivores can elicit evolutionary responses in some traits of non‐prey species. In general, life history traits of unpalatable species may be cryptically adapting to stressful indirect effects where large herbivores are overabundant.     

Trait‐mediated indirect interactions in a marine intertidal system as quantified by functional responses

Studies of trait‐mediated indirect interactions (TMIIs) typically focus on effects higher predators have on per capita consumption by intermediate consumers of a third, basal prey resource. TMIIs are usually evidenced by changes in feeding rates of intermediate consumers and/or differences in densities of this third species. However, understanding and predicting effects of TMIIs on population stability of such basal species requires examination of the type and magnitude of the functional responses exhibited towards them. Here, in a marine intertidal system consisting of a higher‐order fish predator, the shanny Lipophrys pholis, an intermediate predator, the amphipod Echinogammarus marinus, and a basal prey resource, the isopod Jaera nordmanni, we detected TMIIs, demonstrating the importance of habitat complexity in such interactions, by deriving functional responses and exploring consequences for prey population stability. Echinogammarus marinus reacted to fish predator diet cues by reducing activity, a typical anti‐predator response, but did not alter habitat use. Basal prey, Jaera nordmanni, did not respond to fish diet cues with respect to activity, distribution or aggregation behaviour. Echinogammarus marinus exhibited type II functional responses towards J. nordmanni in simple habitat, but type III functional responses in complex habitat. However, while predator cue decreased the magnitude of the type II functional response in simple habitat, it increased the magnitude of the type III functional response in complex habitat. These findings indicate that, in simple habitats, TMIIs may drive down consumption rates within type II responses, however, this interaction may remain de‐stabilising for prey populations. Conversely, in complex habitats, TMIIs may strengthen regulatory influences of intermediate consumers on prey populations, whilst potentially maintaining prey population stability. We thus highlight that TMIIs can have unexpected and complex ramifications throughout communities, but can be unravelled by considering effects on intermediate predator functional response types and magnitudes. Synthesis Higher‐order predators and habitat complexity can influence behaviour of intermediate species, affecting their consumption of prey through trait‐mediated indirect interactions (TMIIs). However, it is not clear how these factors interact to determine prey population stability. Using functional responses (FRs), relating predator consumption to prey density, we detected TMIIs in a marine system. In simple habitats, TMIIs reduced consumption rates, but FRs remained de‐stabilising for prey populations. In complex habitats, TMIIs strengthened prey regulation with population stabilizing FRs. We thus demonstrate that FRs can assess interactions of environmental and biological cues that result in complex and unexpected outcomes for prey populations.     

Indirect interactions in a rice ecosystem: density dependence and the interplay between consumptive and non‐consumptive effects of predators

1. Density‐ and trait‐mediated indirect interactions (DMIIs and TMIIs, respectively) in food chains play crucial roles in community structure and processes. However, factors affecting the relative strength of these interactions are poorly understood, including in widespread and important freshwater rice ecosystems. 2. We studied the strength of DMIIs and TMIIs in a food chain involving a predator (the Reeve’s turtle Chinemys reevesii), its herbivorous prey (the apple snail Pomacea canaliculata) and a plant (rice Oryza sativa) in outdoor containers simulating rice fields. We also evaluated consumptive and non‐consumptive effects of the predator on the snail. We removed a fixed proportion of snails every 2 days to simulate prey consumption and introduced a caged turtle that was fed daily with snails to simulate non‐consumptive effects. 3. Direct consumptive effects increased growth of the remaining snails and their per capita feeding rate. Moreover, consumptive and non‐consumptive effects, and their interaction, affected the proportion of snails buried in the soil. This interaction was presumably because increasing food availability per snail induced their self‐burying behaviour. 4. Both DMIIs and TMIIs affected the number of rice plants remaining, whereas their interaction term was not significant. 5. In summary, density dependence and interactions between consumptive and non‐consumptive effects influenced snail growth and behaviour, respectively. However, no cascading effects of these complicated interactions on rice plants were detected.     

Local adaptation of fish consumers alters primary production through changes in algal community composition and diversity

Ecological research has focused on understanding how changes in consumer abundance affects community structure and ecosystem processes. However, there is increasing evidence that evolutionary changes in consumers can also alter community structure and ecosystem processes. Typically, the effects of consumer phenotype on communities and ecosystem processes are measured as net effects that integrate numerous ecological pathways. Here, we analyze new data from experimental manipulations of Trinidadian guppy Poecilia reticulata presence, density and phenotype to examine how effects on the algal community cause changes in gross‐primary production (GPP). We combine analytical tools borrowed from path analysis with experimental exclosures in mesocosms to separate the ecological and evolutionary effects of guppies into direct and indirect components. We show that the evolutionary effects of guppy phenotype act through different ecological pathways than the effects of guppy presence and density on GPP. As reported in previous studies that used a different measure of algal biomass, adding guppies and doubling their densities decreased algal biovolume through direct effects. In contrast to these previously reported results, exchanging guppy phenotypes that live without predators for phenotypes that live with predators did not affect algal biovolume. Instead, guppies from populations that live with predators increased the diversity of algal species and increased GPP compared to guppies that live without predators. These changes in the algal community were driven primarily by guppy phenotypes that live with predators—algal communities in mesocosms without fish were similar to those with guppies from predator‐free locations, but both were different from mesocosms with guppies from populations that live with predators. Changes in the algal community were driven directly by differences in foraging behavior between the two consumer phenotypes. We reconcile these results with our previous findings, thereby enhancing our understanding of the relationship between ecological and evolutionary processes.     

Multi-predator effects across life-history stages: non-additivity of egg- and larval-stage predation in an African treefrog

The effects of multiple predators on their prey are frequently non‐additive because of interactions among predators. When prey shift habitats through ontogeny, many of their predators cannot interact directly. However, predators that occur in different habitats or feed on different prey stages may still interact through indirect effects mediated by prey traits and density. We conducted an experiment to evaluate the combined effects of arboreal egg‐stage and aquatic larval‐stage predators of the African treefrog, Hyperolius spinigularis. Egg and larval predator effects were non‐additive – more Hyperolius survived both predators than predicted from their independent effects. Egg‐stage predator effects on aquatic larval density and size and age at hatching reduced the effectiveness of larval‐stage predators by 70%. Our results indicate that density‐ and trait‐mediated indirect interactions can act across life‐stages and habitats, resulting in non‐additive multi‐predator effects.     

Aggressive Mimicry by the Characid Fish Erythrinus erythrinus

The young Erythrinus erythrinus mimics the colour pattern of the female of its specific prey, the cyprinodont fish Rivulus agilae. The male Rivulus is attracted by the predator, and, when performing sexual display in front of it, is caught by the tail and swallowed. Various experiments in captivity confirm the constancy of this behavioural pattern. The morphological, ecological and behavioural specializations related to the aggressive mimicry by Erythrinus are described. The colour pattern of the young Erythrinus disappears completely when the fish becomes subadult and leaves the highly specialized habitat of the Rivulus, to settle in deeper water. Comparisons between the present case and other described cases of aggressive mimicry are made. The evolutionary processes involved in aggressive mimicry displayed by Erythrinus are examined: origin of the morphological adaptations in the predator, consequences for the population dynamics in the prey.     

Temperature and prey density jointly influence trophic and non‐trophic interactions in multiple predator communities

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