Prey nutrient composition has different effects on Pardosa wolf spiders with dissimilar life histories

The nutritional composition of prey is known to influence predator life histories, but how the life history strategies of predators affect their susceptibility to nutrient imbalance is less investigated. We used two wolf spider species with different life histories as model predators: Pardosa amentata, which have a fixed annual life cycle, and Pardosa prativaga, which reproduce later and can extend development across 2 years. We fed juvenile spiders of the two species ad libitum diets of one of six Drosophila melanogaster fly types varying in lipid:protein composition during three instars, from the start of the second instar until the fifth instar moult. We then tested for interactions between predator species and prey nutrient composition on several life history parameters. P. amentata completed the three instars faster and grew larger carapaces and heavier body masses than P. prativaga, but the two species responded differently to variation in prey lipid:protein ratio. Duration of the instars increased when feeding on protein-poor prey in P. amentata, but was unaffected by diet in P. prativaga. Likewise, the effect of diet on body composition was more pronounced in P. amentata than in P. prativaga. Prey nutrient composition thus affected the two species differently. During macronutrient imbalance P. amentata appear to prioritize high growth rates while experiencing highly variable body compositions, whereas P. prativaga maintain more constant body compositions and have slower growth. These can be seen as different consequences of a fixed annual and a plastic annual-biennial life cycle.     

Food quality effects on instar‐specific life histories of a holometabolous insect

1. It is a long‐standing challenge to understand how changes in food resources impact consumer life history traits and, in turn, impact how organisms interact with their environment. To characterize food quality effects on life history, most studies follow organisms throughout their life cycle and quantify major life events, such as age at maturity or fecundity. From these studies, we know that food quality generally impacts body size, juvenile development, and life span. Importantly, throughout juvenile development, many organisms develop through several stages of growth that can have different interactions with their environment. For example, some parasitoids typically attack larger instars, whereas larval insect predators typically attack smaller instars. Interestingly, most studies lump all juvenile stages together, which ignores these ecological changes over juvenile development.
2. We combine a cross‐sectional experimental approach with a stage‐structured population model to estimate instar‐specific vital rates in the bean weevil, Callosobruchus maculatus across a food quality gradient. We characterize food quality effects on the bean weevil's life history traits throughout its juvenile ontogeny to test how food quality impacts instar‐specific vital rates.
3. Vital rates differed across food quality treatments within each instar; however, their effect differed with instar. Weevils consuming low‐quality food spent 38%, 37%, and 18% more time, and were 34%, 53%, and 63% smaller than weevils consuming high‐quality food in the second, third, and fourth instars, respectively. Overall, our results show that consuming poor food quality means slower growth, but that food quality effects on vital rates, growth and development are not equal across instars. Differences in life history traits over juvenile ontogeny in response to food quality may impact how organisms interact with their environment, including how susceptible they are to predation, parasitism, and their competitive ability.

     

Effects of prey density, prey instar, and patch size on the development of the predatory mosquito, Toxorhynchites towadensis

Effects of prey density, prey instar, and patch size on the development of the predatory mosquito larva, Toxorhynchites towadensis, were investigated in the laboratory. Survivors of T. towadensis showed different developmental patterns in relation to prey age structure. All predatory larvae in containers with only second instar prey developed into the third instar. However, in several containers with fourth instar prey, mortality of predators was observed. During the third instar, no predatory larva died, but both prey density and prey instar significantly affected the survival of predators during their fourth instar. Large prey size promoted large predator adults, and predatory larvae which grew up in small surface containers responded by developing to large sizes than those in large containers. Larval developmental time of the predators differed in each treatment. During first and second instars, faster predator development was observed in containers with small surface areas and containing young prey individuals. However, when development was enhanced by the presence of old prey individuals, no surface effect was observed. The fastest predator development was observed with prey of mixed instars and high density. This study suggests that a small surface container containing prey of mixed instars and high density is suitable for development of predators.     

Stable isotope analyses document intraguild predation in wolf spiders (Araneae: Lycosidae) and underline beneficial effects of alternative prey and microhabitat structure on intraguild prey survival

Intraguild predation (IGP) is common among generalist predators and an important issue in food web theory, because IGP may destabilise communities by increasing extinction of species. Also, IGP may interfere with the effectiveness of generalist predators as biological control agents. In general, occurrence of IGP in laboratory or field studies is inferred from abundance data or direct observations only. We investigated if tracing stable isotopes allows distinction between different types of predation and confirmation of IGP. Wolf spiders were chosen as model organisms for generalist predators; IGP of third instar A. cuneata on second instar P. palustris was investigated in a laboratory experiment. The availability of alternative prey and the complexity of the microhabitat were manipulated, since both factors are thought to facilitate coexistence of predators.
Stable isotope analysis documented predation of A. cuneata on P. palustris and predation on alternative prey by both juveniles. Both the presence of alternative prey and the availability of shelter reduced mortality of juvenile P. palustris during the first week. During the second week mortality increased in complex structure without alternative prey presumably due to enhanced activity and cannibalism among starving P. palustris . Thus, microhabitat complexity and prey abundance may foster coexistence of wolf spiders in the field.
In conclusion, stable isotope analysis was proven a powerful tool to investigate animal behaviour without direct observation. The method allowed disentangling predator feeding behaviour when more than one type of prey was present.     

Predation ability and non-consumptive effects of Notonecta sellata (Heteroptera: Notonectidae) on immature stages of Culex pipiens (Diptera: Culicidae)

Predators may have multiple effects on prey, including the mortality caused by consumption, but also non-consumptive effects when prey alter their life history traits in the presence of predators. This study aimed to describe the consumption ability and the non-consumptive effects of Notonecta sellata (Heteroptera: Notonectidae) on immature stages of Culex pipiens (Diptera: Culicidae). Results showed that adult N. sellata were capable of preying on all larval instars, although they consumed more individuals of the 2(nd) and 3(rd) instars. Immature mosquitoes raised in the presence of, but without contact with, predators showed a slower development and smaller-sized emerging adults than those raised in the control treatments. Similar survival rates were recorded in the predator and control treatments. The present study suggests that N. sellata adults negatively affect Cx. pipiens populations in two ways: a) by increasing immature stage mortality as a result of direct consumption and extended development times; and b) by reducing their number of offspring, as a result of delayed reproduction and a lower fecundity of adults.     

Weak responses to dietary enrichment in a specialized aphid predator

Some ladybeetles are specialist predators of aphids, coccids or other prey, although they often eat a variety of species from their focal prey taxon. In addition, the diet is often supplemented with alternative prey. How larvae of the aphidophagous Coccinella septempunctata L. utilize a non‐aphid alternative prey (fruit‐fly larvae Drosophila melonogaster Meigen) is compared with adequate (i.e. high‐quality) aphid prey provided alone (monotypic diet) or in mixed diets. The alternative prey are presented either nutrient‐enriched (i.e. raised on dog food supplemented medium) or not (raised on pure medium). Ladybird performance (survival, growth and development) is poor on the pure fly larvae diets, and also reduced when given mixed diets compared with the pure aphid diet. Nutrient enrichment of the fly larvae has no positive effects. The physiological background for the differences in food value, as indicated by performance in life‐history parameters, is a strong pre‐ingestive effect (i.e. reduced consumption of fly larvae compared with aphids) and a post‐ingestive effect (i.e. reduced utilization of assimilated larval fly tissue), whereas the assimilation efficiency of the consumed fly larvae is as high as that of aphids. The results show a physiological trade‐off resulting from prey specialization that reduces the possibility of utilizing alternative prey when the availability of aphids is scarce. Connected with this is a high robustness against variation in prey nutrient diversity and composition; the ladybird shows little positive response to dietary mixing (i.e. neither mixing of adequate aphids, nor of aphids and alternative prey) or to nutrient enrichment of prey. This contrasts with the results from generalist predators (spiders), where similar treatments lead to strong effects on life‐history parameters.     

Nutrient balance affects foraging behaviour of a trap-building predator

Predator foraging may be affected by previous prey capture, but it is unknown how nutrient balance affects foraging behaviour. Here, we use a trap-building predator to test whether nutrients from previous prey captures affect foraging behaviour. We fed orb-weaving spiders (Zygiella x-notata) prey flies of different nutrient composition and in different amounts during their first instar and measured the subsequent frequency of web building and aspects of web architecture. We found that both the likelihood of web building and the number of radii in the web were affected by prey nutrient composition while prey availability affected capture area and mesh height. Our results show that both the balance of nutrients in captured prey and the previous capture rate may affect future foraging behaviour of predators.     

Ontogenetic shifts in the functional response and interference interactions of Rhyacophila dorsalis larvae (Trichoptera)

1. Ontogenetic shifts in predator behaviour can affect the assessment of food‐web structure and the development of predator–prey models. Therefore, it is important to establish if the functional response and interference interactions differ between life‐stages. These hypotheses were tested by (i) comparing the functional response of second, third, fourth and fifth larval instars of Rhyacophila dorsalis, using three stream tanks with one Rhyacophila larva per tank and one of 10 prey densities between 20 and 200 larvae of Chironomus sp.; (ii) using other experiments to assess interference within instars (two to five larvae of the same instar per tank), and between pairs of different instars (one, two or three larvae per instar; total predator densities of two, four or six larvae per tank). 2. The first hypothesis was supported. The number of prey eaten by each instar increased with prey density, the relationship being described by a type II model. The curvilinear response was stronger for fourth and fifth instars than for second and third instars. Mean handling time did not change significantly with prey density, and increased with decreasing instar number from 169 s for fifth instars to 200 s for second instars. Attack rate decreased progressively with decreasing instar number. Handling time varied considerably for each predator–prey encounter, but was normally distributed for each predator instar. Variations in attack rate and handling time were related to differences in activity between instars, fourth and fifth instars being more active and aggressive than second and third instars, and having a higher food intake. 3. The second hypothesis was partially supported. In the interference experiments between larvae of the same instar or different instars, mean handling time did not change significantly with increasing predator density, and attack rate did not change for second and third instars but decreased curvilinearly for fourth and fifth instars. Interference between some instars could not be studied because insufficient second instars were available at the same time as fourth and fifth instars, and most third instars were eaten by fourth and fifth instars in the experiments. Prey capture always decreased with decreasing attack rate. Therefore, interference reduced prey consumption in fourth and fifth instars, but not in second and third instars. The varying feeding responses of different instars should be taken into account when assessing their role in predator–prey relationships in the field.     

A laboratory investigation of mosquito larval predation by Toxorhynchites moctezuma on Aedes aegypti

1. Functional responses of predatory Toxorhynchites moctezuma (Dyar & Knab) larvae feeding on Aedes aegypti (L.) larvae (Diptera: Culicidae) were found to be type II of Holling (1959) and Rogers (1972). 2. Estimates of searching rate were generally higher for later instar predators. The search rate of second instar predators declined as prey instar increased, but fourth instar Tx. moctezuma had the highest search rate for second instar Ae. aegypti. 3. Prey handling times were higher for early instar predators and late instar prey. 4. When presented with mixtures of two instars of Ae. aegypti, second instar Tx. moctezuma showed frequency independent selectivity for the early instars, whereas fourth instar predators showed frequency independent selectivity for the late instars of Ae. aegypti. There was no evidence of frequency dependent predation. Preferences appeared to be transitive. 5. Extended random predator equations, using parameters derived from the functional responses, did not adequately describe the outcome of predation in the prey mixture experiment, even when the possibility of optimal switching behaviour was accounted for.     

Voracity and prey preference of insidious flower bug (hemiptera: anthocoridae) for immature stages of soybean aphid (hemiptera: aphididae) and soybean thrips (thysanoptera: thripidae)

The generalist predator, Orius insidiosus (Say), is an important natural enemy of the soybean aphid, Aphis glycines Matsumura. Soybean thrips, Neohydatothrips variabilis (Beach), serve as an important prey resource for O. insidiosus in soybeans, sustaining the predator's population before the arrival of the soybean aphid. Although generalist predators can forage on a broad range of prey, they may show distinct preferences for particular prey, attacking prey at levels disproportionate to their relative numbers. To assess the preference of O. insidiosus for soybean aphid and soybean thrips, attack rates of nymphal and adult O. insidiosus were measured in the laboratory. For both adults and nymphs, the number of prey attacked increased as more prey were provided. For nymphs, the total number of prey attacked increased as the predator matured. In general, the number of prey attacked by adult predators was relatively constant as the predator aged. Both O. insidiosus nymphs and adults displayed a preference for soybean thrips, by disproportionately attacking soybean thrips over soybean aphid regardless of the relative densities of the two prey. We discuss implications of this preference on O. insidiosus life history characteristics and the potential impact on O. insidiosus-prey dynamics in the field.     

Consequences of divergent egg phenology to predation and coexistence in two sympatric,congeneric mantids (Orthoptera: Mantidae)

Summary Two sympatric, congeneric mantid species, Tenodera sinensis (Saussure) and t. angustipennis (Saussure) exhibit intrinsically different temporal patterns of egg hatch in the spring. Tenodera sinensis hatches first, and begins to enter second instar by the time T. angustipennis starts to hatch. As a result, there is asynchrony in developmental times such that several different instars may be present during the first crucial month of life when mantid population densities are high and prey availability is low. The body length ratio of larger to smaller nymphs is most commonly 1.37, in favor of T. sinensis. This character divergence suggests allochronic niche separation which may alleviate competition for prey. However this same ratio appears to be the threshold for both interspecific predation and cannibalism among these bitrophic generalist predators. Therefore T. sinensis, which is nearly always larger, may have a selective advantage by utilizing its smaller congener and later-hatching conspecifics as prey in a food limited environment.     

Predation, individual variability and vertebrate population dynamics

Both predation and individual variation in life history traits influence population dynamics. Recent results from laboratory predator–prey systems suggest that differences between individuals can also influence predator–prey dynamics when different genotypes experience different predation-associated mortalities. Despite the growing number of studies in this field, there is no synthesis identifying the overall importance of the interactions between predation and individual heterogeneity and their role in shaping the dynamics of free-ranging populations of vertebrates. We aim to fill this gap with a review that examines how individual variability in prey susceptibility, in predation costs, in predator selectivity, and in predatory performance, might influence prey population dynamics. Based on this review, it is clear that (1) predation risk and costs experienced by free-ranging prey are associated with their phenotypic attributes, (2) many generalist predator populations consist of individual specialists with part of the specialization associated with their phenotypes, and (3) a complete understanding of the population dynamic consequences of predation may require information on individual variability in prey selection and prey vulnerability. Altogether, this work (1) highlights the importance of maintaining long-term, detailed studies of individuals of both predators and prey in contrasting ecological conditions, and (2) advocates for a better use of available information to account for interactive effects between predators and their prey when modelling prey population dynamics.     

Combined effects of allelochemicals, prey availability, and supplemental plant material on growth of a generalist insect predator

We examined the effects of the presence of plant allelochemicals in prey diet, prey availability and supplemental plant material on the growth of the generalist predator Podisus maculiventris (Hemiptera: Pentatomidae). We tested two different nymphal stages of this predator. Third to fourth instar nymphs and fifth instar nymphs were fed a diet of prey (Manduca sexta larvae, Lepidoptera: Sphingidae) without allelochemicals in their diet or prey fed maximal levels of allelochemicals (tomatine, rutin and chlorogenic acid) found in their host plant (Lycopersicon esculentum). The nymphs were fed prey ad libitum, once every three days, or once every five days. They were given either no supplemental plant material or a 2 cm slice of green bean pod (Phaseolus vulgaris). We also conducted another experiment with fifth instar nymphs using the same conditions, except that mean levels of allelochemicals found in the host plant were fed to prey instead of maximal levels and the prey were provided either once a day or once every five days. For all experiments, prey scarcity depressed developmental rate, weight gain and relative growth rate. Overall, there was no negative effect of allelochemicals in the diet of the prey on these variables when predators were supplied with an excess of prey, but allelochemicals in the prey diet negatively affected these predators when prey were scarce. The addition of plant material to the diet of third to fourth instar nymphs did not have any effect on developmental rate, final dry weight, or relative growth rate. However, for fifth instar nymphs, the addition of plant material negatively affected these variables. Thus, the addition of plant material to the diet of the nymphs did not alleviate the negative effects of prey scarcity or allelochemicals in prey diet.     

Control of aphids on wheat by generalist predators: effects of predator density and the presence of alternative prey

There is evidence for both positive and negative effects of generalist predators on pest populations and the various reasons for these contrasting observations are under debate. We studied the influence of a generalist predator, Pardosa lugubris (Walckenaer) (Araneae: Lycosidae), on an aphid pest species, Rhopalosiphum padi (L.) (Hemiptera: Aphididae; low food quality for the spider), and its host plant wheat, Triticum spec. (Poaceae). We focused on the role of spider density and the availability of alternative prey, Drosophila melanogaster Meigen (Diptera: Drosophilidae; high food quality). The presence of spiders significantly affected plant performance and aphid biomass. Alternative prey and spider density strongly interacted in affecting aphids and plants. High spider density significantly improved plant performance but also at low spider density plants benefited from spiders especially in the presence of alternative prey. The results suggest that generalist arthropod predators may successfully reduce plant damage by herbivores. However, their ability to control prey populations varies with predator nutrition, the control of low-quality prey being enhanced if alternative higher-quality prey is available.     

The demographic and life‐history costs of fear: Trait‐mediated effects of threat of predation on Aedes triseriatus

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Predators alter the scaling of diversity in prey metacommunities

Although predator effects on the number of locally coexisting species are well understood, there are few formal predictions of how these local predator effects influence patterns of prey diversity at larger spatial scales. Building on the theory of island biogeography, we develop a simple model that describes how predators can alter the scaling of diversity in prey metacommunities and compares the effects of generalist and specialist predators on regional prey diversity. Generalist predators, which consume prey randomly with respect to species identity, are predicted to reduce α‐diversity and increase β‐diversity thereby maintaining regional diversity (γ‐diversity). Alternatively, specialist predators, which filter out prey species intolerant of predators, are predicted to reduce bothα‐diversity andβ‐diversity by causing the same prey species to be extirpated in each locality, resulting in regional prey species extinctions and lower γ‐diversity. These distinct effects of generalist and specialist predators on prey diversity at different spatial scales are uniquely shaped by the extent of predation within those metacommunities. Overall, our model results make general predictions for how different types of predators can differentially affect prey diversity across spatial scales, allowing a more complete understanding of the possible implications of predator eradications or introductions for biodiversity.     

Food selectivity and diel vertical distribution of Chaoborus edulis (Diptera, Chaoboridae) in Lake Malawi

1. Diel diet and vertical distribution patterns of the larval instars of Chaoborus edulis were studied in deep water near the central part of Lake Malawi, Africa.
2. First instar larvae contained very little food in their crops and probably depended on reserves from the egg. Second, third and fourth instars fed on zooplankton and were size-selective in their feeding. The mean size of prey eaten by the three instars was significantly different from each other, with larger instars feeding on larger prey. Smallest available prey was selected against and the upper size of prey was probably constrained by larval gape. Nauplii were not found in any of several thousand larvae examined. Phytoplankton did not form a significant part of the diet.
3. There was a progressive and related increase in diel periodicity in feeding and vertical migrations of successive instar stages. Fourth instars migrated particularly large distances. Such migrations removed them from their zooplankton food supply but avoided predators. A refuge from predators is probably found in or near the permanent zero oxygen boundary, at depths greater than 200 m.     

Snail predation by larvae of Sepedon scapularis Adams (Diptera: Sciomyzidae), a potential biocontrol agent of snail intermediate hosts of schistosomiasis in South Africa

Malacophagous larvae of the fly Sepedon scapularis Adams were shown experimentally to be effective predators of three species of aquatic pulmonate snails tested as prey: Bulinus africanus (Krauss) an important intermediate host of Schistosoma haematobium (Bilharz), Bulinus tropicus (Krauss) and the invasive species Physa acuta Draparnaud. Survival of S. scapularis larvae from instar to instar was negatively affected by the size of prey snails, since larvae tended to be asphyxiated by the mucous secretions of the snails, or by the larval hydrofuge hairs becoming entangled in snail faeces. In experiments to test the choice of S.scapularis larvae for different species and sizes of snails, B.africanus was significantly disfavoured compared with the other two snail species. Small snails (< 3 mm) were more frequently killed by all three larval instars of S.scapularis. First instars killed few, if any, large snails (> 7 mm), whereas second and third instars preyed effectively on all sizes of snails. Third instars killed significantly more snails than younger instars. For larvae offered only one species of snail as prey, the mean total number of snails killed per larva during its entire development was 49 B.africanus, 45 B.tropicus or 34 P.acuta. It is concluded that S.scapularis is potentially useful as a biological control agent for use against indigenous Bulinus and exotic Physa snails.     

Cross‐ecosystem disturbance influences on the life history and population size structure of a riparian predator

Cross‐ecosystem transfers of resources could alter the life history traits of consumers in adjacent systems by changing the nature and availability of prey. However, large‐scale influences, such as natural disturbances, that control the magnitude of prey subsidies are likely to modify these effects. To investigate impacts of cross‐ecosystem subsidies on the life history traits of a riparian predator we measured the size, sex and condition of riparian fishing spiders (Dolomedes aquaticus) across a gradient of flooding frequency and intensity. These spiders rely on adult aquatic insects for a large proportion of their diet and previous research demonstrated that increased flooding decreased the abundance of aquatic insect prey. In this study, laboratory experiments indicated that increased prey availability hastened the first moult of the spiders after winter and decreased the propensity for cannibalistic interactions of individuals of the same size. However, despite the likely positive influences of increased food supply, in the field the highest abundance and proportion of large, potentially reproductive females occurred at the most flood‐prone rivers, where aquatic prey availability was the lowest. It is likely that other factors modified by the disturbance regime, such as habitat availability, flood‐related mortality and intra‐specific interaction rates, altered the influence of cross‐ecosystem subsidies on the life history traits of these spiders. Thus, our results indicate that disturbance‐related effects can flow across ecosystem boundaries and alter the life history traits of predators relying on allochthonous resources.     

Effects of Bt-toxin Cry1Ac on Propylaea japonica Thunberg (Col., Coccinellidae) by feeding on Bt-treated Bt-resistant Helicoverpa armigera (Hübner) (Lep., Noctuidae) larvae

Abstract:  Propylaea japonica is an important predatory insect of common cotton pests. To assess the ecological effects of transgenic Bt cotton, expressing Cry1Ac toxin, on this predator, we examined the life history parameters of P. japonica for two generations by feeding them with Bt-resistant Helicoverpa armigera . After ingesting Bt-treated Bt-resistant H. armigera larvae in the third and fourth instar, the body mass and body length of adult P. japonica decreased, a combined effect of poor prey quality and Cry1Ac Bt-toxin may account for these effects. However, larval survivorship and development in these two instars, pupal mortality, fecundity and adult longevity of P. japonica were not affected in both the generations. These results suggest that ingesting Bt-toxin Cry1Ac-treated pests in advanced larval stage might have no significant effect on the fitness of predator P. japonica .