Predator-induced plasticity in guppy (<Emphasis Type="Italic">Poecilia reticulata</Emphasis>) life history traits

A number of invertebrates show predator-induced plasticity in life-history and morphological traits that are considered adaptive. Evidence is accumulating that vertebrates may also adjust their life-history traits in response to predators; however, some of the patterns of plasticity, which appear to be an adaptive response specifically to the risk of size-selective predation, may instead result from reduced foraging in response to predator presence. Here, we describe a study of predator-induced plasticity in guppies (Poecilia reticulata). We have predicted that the plastic response to cues from a small, gape-limited, natural predator of guppies, the killlifish (Rivulus hartii), would be the opposite of that caused by reduced food intake. We have found that male guppies increased their size at maturity, both length and mass, in response to the non-lethal presence of this predator. This pattern of plasticity is the opposite of that observed in response to reduced food intake, where male guppies reduce size at maturity. The increase in size at maturity that we observed would likely reduce predation on adult male guppies by this native predator because it is gape-limited and can only eat juvenile and small adult guppies. This size advantage would be important especially because male guppies grow very little after maturity. Therefore, the pattern of plasticity that we observed is likely adaptive. In contrast, female guppies showed no significant response in size at first parturition to the experimental manipulation; however, we did find evidence suggesting that females may produce more, smaller offspring in response to cues from this predator.     

Life-history consequences of predation for a subantarctic beetle: evaluating the contribution of direct and indirect effects

1. Recently, a small predatory beetle, Trechisibus antarcticus (Carabidae), was accidentally introduced onto the island of South Georgia, sub-Antarctic.
2. From the presumed site of introduction the beetle is invading the coastal lowland area, building up high densities locally in the tussock-forming grass Parodiochloa flabellata .
3. In the coastal area the endemic detritivorous/herbivorous beetle Hydromedion sparsutum (Perimylopidae) is common, especially in and beneath the tussocks.
4. The first three, out of six, larval instars of H. sparsutum are easily taken prey by the carabid.
5. In sites colonized by the carabid, total abundance and the ratio between larval and adult numbers of H. sparsutum are far lower, and its adult body size clearly larger, than in comparable sites where the carabid is absent.
6. Two hypotheses are proposed for explaining the increase in adult body size of H. sparsutum : (i) the increase is a direct effect of predation: selection by the predator favours large hatchlings and/or larvae with a high growth rate; and (ii) the increase is an indirect effect of predation: by lowering the density of H. sparsutum , predation has increased its per capita food supply, enabling a higher growth rate and a larger adult body size.
7. A food addition experiment in a carabid-free site showed availability of high quality food to be insufficient for sustaining the initial larval population.
8. In the laboratory, females from the predator-infested sites produced larger eggs and hatchlings than females from the carabid-free sites, but mass specific growth rates of the larvae were not higher.
9. Field and laboratory data give stronger support to the food hypothesis than to the size selectivity hypothesis.     

Facilitation of fisheries by natural predators depends on life history of shared prey

Predators commonly share prey with human exploiters, intuitively suggesting that there is an inherent human–predator conflict through competition for prey. Here we studied the effects of fishing and predation mortality on biomass distributions and yields of shared prey using a size‐structured model of competing populations, describing the life histories of Baltic Sea sprat and herring. Whereas both species responded in a similar fashion to increased fishing mortality, with decreasing juvenile and adult biomasses, we found that responses to predation mortality differed between species. Sprat only display weak compensatory responses with increasing predation mortality, while over a substantial range of mortalities there was a strong increase in adult (and total) herring biomass, i.e. overcompensation. The observed biomass overcompensation results from relaxed intraspecific competition as predation mortality increased, allowing for faster individual growth rates that in turn lead to a change in population composition (juvenile:adult biomass ratio). Our results suggest that the potential for biomass overcompensation is higher for species exhibiting substantial growth after maturation. Differences in size‐selectivity of predators and fishing mortality resulted in a positive effect of predation mortality on fisheries yields, which can be explained by an overcompensatory response in adult herring biomass. Thus, somewhat counter intuitive, our results suggest that fishermen, depending on prey life history, may actually benefit from allowing for a higher abundance of predators, despite competing for shared prey.     

Effect of resource gradient on age and size at maturity and their influence on early‐life fecundity in the predatory Asian lady beetle,Harmonia axyridis

Variation in food availability impacts the performance of insects in terms of their size and age to maturity and fecundity. Age at maturity determines how quickly individuals in a population can start to reproduce and how much they can reproduce. Results from studies on various insect species show that food availability influences the size and fecundity of adult females. It is predicted that under poor growth conditions, variation in size is low, but variation in age at maturity is considerable. This prediction was examined in a widely distributed lady beetle species, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), a predator of aphids and coccids. Using a food gradient from low to high aphid prey density, performance of females that were reared on excess food was recorded for pre‐reproductive duration, size at reproductive maturity, number of aphids consumed, and fecundity in the first 10 days of their reproductive period. Results suggested that female H. axyridis that were reared on surplus food when kept at low prey density (poor growth condition) took, on average, three times longer to attain maturity and produced, on average, 14 times fewer eggs than females that were also reared on surplus food, but kept at high prey density (good growth condition). Females performed best at a prey density of 30 aphids per female per 150 cm². Results suggested that the current food availability significantly influenced the age and size of females at maturity and their fecundity. Age and size at maturity of female lady beetles showed non‐linear responses to prey density as well as the occurrence of a minimum size of females, below which H. axyridis females fail to mature. The steep slope recorded at lower prey densities suggests relatively high variation in age at maturity but low variation in size.     

Coexistence of predator and prey in intraguild predation systems with ontogenetic niche shifts

In basic intraguild predation (IGP) systems, predators and prey also compete for a shared resource. Theory predicts that persistence of these systems is possible when intraguild prey is superior in competition and productivity is not too high. IGP often results from ontogenetic niche shifts, in which the diet of intraguild predators changes as a result of growth in body size (life-history omnivory). As a juvenile, a life-history omnivore competes with the species that becomes its prey later in life. Competition can hence limit growth of young predators, while adult predators can suppress consumers and therewith neutralize negative effects of competition. We formulate and analyze a stage-structured model that captures both basic IGP and life-history omnivory. The model predicts increasing coexistence of predators and consumers when resource use of stage-structured predators becomes more stage specific. This coexistence depends on adult predators requiring consumer biomass for reproduction and is less likely when consumers outcompete juvenile predators, in contrast to basic IGP. Therefore, coexistence occurs when predation structures the community and competition is negligible. Consequently, equilibrium patterns over productivity resemble those of three-species food chains. Life-history omnivory thus provides a mechanism that allows intraguild predators and prey to coexist over a wide range of resource productivity.     

What keeps insects small?—Size dependent predation on two species of butterfly larvae

Insect size usually increases greatly in the latter stages of development, while reproductive value increases strongly with adult size. Mechanisms that can balance the benefits associated with increased growth are poorly understood, raising the question: what keeps insects from becoming larger? If predation risk was to increase with juvenile size, it would make an extension of development very risky, favouring smaller final sizes. But field measures of juvenile mortality seldom show any general patterns of size dependence. We here therefore try to estimate a mechanistic relationship between juvenile size and predation risk by exposing the larvae of two closely related butterflies to a generalist invertebrate predator in a laboratory experiment. Predation risk increased with larval size but was not affected by the species-specific growth rate differences. These results indicate that predation risk may increase with the size of the juvenile even when predators are relatively small. By basing a model simulation on our data we also show that size dependent predation of the kind found in this study has potential to stabilise selection on body size in these species. Thus, these findings suggest that more detailed studies of the size dependence of predation risk on juvenile instars will increase the understanding of what it is that keeps insects small.     

Complementary predation on metamorphosing species promotes stability in predator–prey systems

Functionally redundant predation and functionally complementary predation are both widespread phenomena in nature. Functional complementary predation can be found, for example, when predators feed on different life stages of their prey, while functional redundant predation occurs when different predators feed on all life stages of a shared prey. Both phenomena are common in nature, and the extent of differential life-stage predation depends mostly on prey life history; complementary predation is expected to be more common on metamorphosing prey species, while redundant predation is thought to be higher on non-metamorphosing species. We used an ordinary differential equation model to explore the effect of varying degree of complementary and redundant predation on the dynamic properties of a system with two predators that feed on an age-structured prey. Our main finding was that predation on one stage (adult or juvenile) resulted in a more stable system (i.e., it is stable for a wider range of parameters) compared to when the two predators mix the two prey developmental stages in their diet. Our results demonstrate that predator–prey dynamics depends strongly on predators' functionality when predator species richness is fixed. Results also suggest that systems with metamorphosing prey are expected to be more diverse compared to systems with non-metamorphosing prey.     

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.     

Predator induced life-history shifts in a freshwater cladoceran

Summary Life-history theory predicts that maturity and resource allocation patterns are highly sensitive to selective predation. Under reduced adult survival, selection will favour genotypes capable of reproducing earlier, at a smaller size and with a higher reproductive effort. When exposed to water that previously held fish, (size selective predators which prefer larger Daphnia), individuals of Daphnia hyalina reproduced earlier, at a smaller size and had a higher reproductive investment. Hence the prey was able to switch its life history pattern in order to become less susceptible to predation by a specific predator. The cue that evokes the prey response is a chemical released by the predator.     

Cooccurrence of prey species alters the impact of predators on prey performance through multiple mechanisms

When prey are differentially affected by intra and interspecific competition, the cooccurrence of multiple prey species alters the per capita availability of food for a particular prey species which could alter how prey respond to the threat of predation, and hence the overall‐effect of predators. We conducted an experiment to examine the extent to which the nonconsumptive and overall effect of predatory water bugs on snail and tadpole traits (performance and morphology) depended on whether tadpoles and snails cooccurred. Tadpoles and snails differed in their relative susceptibility to intraspecific and interspecific competition, and predators affected both prey species via consumptive and nonconsumptive mechanisms. Furthermore, the overall effect of predators often depended on whether another prey species was present. The reasoning for why the overall effect of predators depended on whether prey species cooccurred, however, differed for each of the response variables. Predators affected snail body growth via nonconsumptive mechanisms, but the change in the overall effect of predators on snail body growth was attributable to how snails responded to competition in the absence of predators, rather than a change in how snails responded to the threat of predation. Predators did not affect tadpole body growth via nonconsumptive mechanisms, but the greater vulnerability of competitively superior prey (snails) to predators increased the strength of consumptive mechanisms (and hence the overall effect) through which predators affected tadpole growth. Predators affected tadpole morphology via nonconsumptive mechanisms, but the greater propensity for predators to kill competitively superior prey (snails) enhanced the ability of tadpoles to alter their morphology in response to the threat of predation by creating an environment where tadpoles had a higher per capita supply of food available to invest in the development of morphological defenses. Our work indicates that the mechanisms through which predators affect prey depends on the other members of the community.     

Variation in growth and age at maturity in bluegill sunfish: genetic or environmental effects?

Populations of bluegill sunfish Lepomis macrochirus , experiencing heavy juvenile predation, showed increased growth rates and increased age and size at maturity relative to populations experiencing decreased predation on juveniles but increased predation on adults. This study examined bluegills experimentally from both types of populations and a cross between them in a common environment to determine if variation in growth and age at maturity is genetically or environmentally induced. Two factorial experiments, varying strain of bluegills and resource availability, were used to evaluate differences in growth rate. One experiment, varying strain of bluegills, was used to assess differences in age at maturity. Growth was strongly influenced by resource level, but growth rate did not vary among populations. Nearly all bluegills in each population matured at 1 year of age in a common environment. Thus, variation observed in source populations must be mostly attributable to differences in the environment between populations. At least three factors could potentially cause differences in growth and age at maturity: (1) variation in resource availability; (2) variation in demographic structure; and (3) variation in size-specific mortality rates caused by differences in predator abundance between populations. Observed patterns of variation between populations are best explained by effects of differences in predator populations.     

Non-lethal effects of predators on prey growth rates depend on prey density and nutrient additions

A number of studies show that predators can depress prey growth rates by inducing reductions in foraging activity, but the size of this non-lethal effect is quite variable. Here I investigate how prey density and resource productivity may alter the extent to which predators depress the growth rates of their prey. Theory predicts that when resources are overgrazed, an increase in predation risk will have little net effect on individual food intake because the decline in foraging effort will be offset by an increase in resource level. Thus, the non-lethal effects of predators on prey growth rates should depend upon prey density and resource productivity in a predictable manner, with the growth penalty imposed by predators being strongest when resources are undergrazed and weakest when resources are overgrazed. I tested this hypothesis by manipulating predation risk, prey density, and nutrient additions in a mesocosm experiment with the pulmonate snail Helisoma trivolvis . Refuge use by snails was 45% higher in the presence of caged crayfish than in their absence. Snail growth rates were reduced, on average, by 24% in the presence of caged crayfish. However, the magnitude of the growth penalty exacted by crayfish depended on snail density and nutrient additions. When snails were stocked at high density and nutrient additions were low, growth suppression was just 2.6%. At the other extreme, when snails were at low density and nutrient additions were high, growth suppression was 44.6%. Thus, the non-lethal effects of predators on prey growth depend on environmental context, illustrating an important link between individual traits and system-level properties.     

The Shrimp Caridina nilotica in Lake Victoria (East Africa), Before and After the Nile Perch Increase

The shrimp Caridina nilotica is a major prey of the introduced Nile perch in Lake Victoria. In spite of heavy predation, the density of shrimps increased after the Nile perch boom and the concomitant disappearance of the haplochromine cichlids. In the same period, the mean size of gravid shrimps and the size at first maturity declined. This seems to indicate an increased predation pressure on adult shrimps. Before the Nile perch upsurge, specialised shrimp eaters and piscivores, among the haplochromine cichlids, only took adult shrimps, whereas we assume that most haplochromines used to include juvenile shrimps into their diet. Another important predator on adult shrimps was Bagrus docmak. The combined density of predators on adult shrimps in the pre-Nile perch era was estimated at 10 kg ha⁻¹ and the potential predators on juveniles were estimated at 170 kg ha⁻¹. After the Nile perch upsurge, only Nile perch up to 10 cm TL and Rastrineobola argentea fed on juvenile shrimps (ca. 36 kg ha⁻¹) and Nile perch from 10 to 50 cm TL (ca. 13 kg ha⁻¹) fed on adults. These rough estimates of the biomass of predators on shrimps before and after the Nile perch upsurge indicate a reduced predation pressure on juvenile shrimps. The disappearance of the haplochromines may have released competition with small Nile perch for juvenile shrimps, thus enhancing the recruitment of Nile perch.     

Early exposure to nonlethal predation risk by size-selective predators increases somatic growth and decreases size at adulthood in three-spined sticklebacks

Predation has an important influence on life history traits in many organisms, especially when they are young. When cues of trout were present, juvenile sticklebacks grew faster. The increase in body size as a result of exposure to cues of predators was adaptive because larger individuals were more likely to survive predation. However, sticklebacks that had been exposed to cues of predators were smaller at adulthood. This result is consistent with some life history theory. However, these results prompt an alternative hypothesis, which is that the decreased size at adulthood reflects a deferred cost of early rapid growth. Compared to males, females were more likely to survive predation, but female size at adulthood was more affected by cues of predators than male size at adulthood, suggesting that size at adulthood might be more important to male fitness than to female fitness.     

Subtle top‐down control of a freshwater meiofaunal assemblage by juvenile fish

1. Top‐down control of prey assemblages by fish predators has been clearly demonstrated in lakes (for zooplankton prey) and rivers (for macroinvertebrate prey). Fish predation can have a significant impact on the body size of prey assemblages; often large‐bodied prey are reduced in abundance, and indirect facilitation of small‐bodied prey occurs potentially initiating a trophic cascade. 2. Benthic communities in aquatic ecosystems also include a numerous and functionally important meiofaunal‐sized component, but in freshwaters the impact of fish predation on meiofaunal assemblages is unknown. We used a laboratory microcosm study to explore the impact of juvenile fish predation on the abundance and size structure of a riverine meiofaunal assemblage. 3. The presence of fish in our microcosms had no significant effect on overall meiofaunal (temporary and permanent) abundance. However, for the Copepoda, we found the first evidence of top‐down control of freshwater meiofaunal assemblages; in microcosms with juvenile fish, the abundance of large‐bodied Copepoda was significantly reduced, whereas small‐bodied Copepoda were significantly more abundant suggesting indirect facilitation. 4. We conclude that predation by juvenile fish can alter the structure of freshwater meiofaunal assemblages, although we do not yet know whether these relatively subtle changes are overwhelmed by large‐scale events such as flow disturbances.     

Life-history responses of a mayfly to seasonal constraints and predation risk

Abstract. 1. An experiment was conducted in the laboratory to examine the effects of photoperiod and predation risk on life-history variation in the mayfly Ephemerella subvaria .
2. Both photoperiod and predation risk affected age at maturity significantly but neither factor affected size at maturity. Mayflies perceiving themselves to be late in the growing season matured in fewer days than those perceiving themselves to be early in the growing season. The presence of predators delayed mayfly maturity significantly.
3. These results suggest that the large variation in life-history traits observed in aquatic insects may be attributed partially to seasonality but that other biotic and abiotic factors may also underlie variation in these traits.     

Predation cues rather than resource availability promote cryptic behaviour in a habitat-forming sea urchin

It is well known that predators often influence the foraging behaviour of prey through the so-called “fear effect”. However, it is also possible that predators could change prey behaviour indirectly by altering the prey’s food supply through a trophic cascade. The predator–sea urchin–kelp trophic cascade is widely assumed to be driven by the removal of sea urchins by predators, but changes in sea urchin behaviour in response to predators or increased food availability could also play an important role. We tested whether increased crevice occupancy by herbivorous sea urchins in the presence of abundant predatory fishes and lobsters is a response to the increased risk of predation, or an indirect response to higher kelp abundances. Inside two New Zealand marine reserves with abundant predators and kelp, individuals of the sea urchin Evechinus chloroticus were rarer and remained cryptic (i.e. found in crevices) to larger sizes than on adjacent fished coasts where predators and kelp are rare. In a mesocosm experiment, cryptic behaviour was induced by simulated predation (the addition of crushed conspecifics), but the addition of food in the form of drift kelp did not induce cryptic behaviour. These findings demonstrate that the ‘fear’ of predators is more important than food availability in promoting sea urchin cryptic behaviour and suggest that both density- and behaviourally mediated interactions are important in the predator–sea urchin–kelp trophic cascade.     

Seasonally varying marine influences on the coastal ecosystem detected through molecular gut analysis

Terrestrial predators on marine shores benefit from the inflow of organisms and matter from the marine ecosystem, often causing very high predator densities and indirectly affecting the abundance of other prey species on shores. This indirect effect may be particularly strong if predators shift diets between seasons. We therefore quantified the seasonal variation in diet of two wolf spider species that dominate the shoreline predator community, using molecular gut content analyses with general primers to detect the full prey range. Across the season, spider diets changed, with predominantly terrestrial prey from May until July and predominantly marine prey (mainly chironomids) from August until October. This pattern coincided with a change in the spider age and size structure, and prey abundance data and resource selection analyses suggest that the higher consumption of chironomids during autumn is due to an ontogenetic diet shift rather than to variation in prey abundance. The analyses suggested that small dipterans with a weak flight capacity, such as Chironomidae, Sphaeroceridae, Scatopsidae and Ephydridae, were overrepresented in the gut of small juvenile spiders during autumn, whereas larger, more robust prey, such as Lepidoptera, Anthomyidae and Dolichopodidae, were overrepresented in the diet of adult spiders during spring. The effect of the inflow may be that the survival and growth of juvenile spiders is higher in areas with high chironomid abundances, leading to higher densities of adult spiders and higher predation rates on the terrestrial prey next spring.     

Safety in numbers for secondary prey populations: an experimental test using egg predation by small mammals in New Zealand

New Zealand's native avifauna is threatened by introduced mammalian predators. Native species are often not the primary prey of these predators, which depend on introduced mice and rabbits as their primary food source. Theoretical models predict that predation risk for a subsidiary, or "secondary" prey species is inversely proportional to its population size. This prediction was tested by a quasi-natural experiment in which four different sized prey "colonies" were constructed at four existing sooty shearwater breeding sites. Domestic hens' eggs were placed in shearwater burrows immediately following the shearwater breeding season and egg predation rates monitored at five, ten and fifteen days. Treatments were switched between sites and the experiment run for a second time after a two-week stand-down period. The net effect of increasing colony size was to lower individual risk of predation. The larger number of individuals present served to effectively "buffer," or dilute, per-capita predation risk from predators whose numbers are fixed by extraneous factors: chiefly the abundance of their primary prey. Although eggs were removed more slowly from smaller colonies than from larger ones, each loss had a greater per-capita effect on individual mortality risk. The inverse density dependent relationship found between colony size and predation risk implies that predator population dynamics are largely independent of secondary prey numbers. Abundant introduced predators can therefore easily drive a small secondary prey population to extinction. Control of primary prey populations may be an important management tool in these circumstances.     

Optimal reproductive strategies in age-structured populations of zooplankton

SUMMARY. We describe a model of zooplankton population dynamics that accounts for differences in mortality and physiology among animals of different ages or sizes. The model follows changes in numbers of individuals and changes in individual and egg biomass through time and it expresses mortality and net assimilation as functions of animal size.
We investigated the effect of egg size, age at first reproduction, and size at first reproduction on the per capita growth rates of populations growing under different conditions. In the absence of predation or when exposed to vertebrate predators that prefer large prey, populations achieve maximum growth rates when animals hatch from small eggs and reach maturity quickly at small sizes. Populations exposed to invertebrate predators that concentrate on small animals may increase r in two different ways. One way is for animals to increase juvenile survivorship by hatching from large eggs and by shortening the juvenile period. An alternative strategy is for animals to hatch from small eggs and to postpone maturity until they grow beyond the range of sizes available to their predators. Certain life history strategies maximize r if animals continue to grow after they reach maturity. By growing larger, non-primiparous females are able to hatch larger clutches and thereby increase the overall rate of population growth.
The model analysis shows how to assess age-dependent mortality rates from field data. The net rate of population increase and the age distribution of eggs together provide specific, quantitative information about mortality.