Land cover diversity increases predator aggregation and consumption of prey

A lower diversity of land cover types is purported to decrease arthropod diversity in agroecosystems and is dependent on patterns of land use and fragmentation. Ants, important providers of ecosystem services such as biological control, are susceptible to landscape‐level changes. We determined the relationships between land cover diversity and fragmentation on the within‐field spatial associations of ants to pests and resulting predation events by combining mapping and molecular tools. Increased land cover diversity and decreased fragmentation increased ant abundance, spatial association to pests and predation. Land cover diversity and fragmentation were more explanatory than land cover types. Even so, specific land cover types, such as deciduous forest, influenced ant and pest diversity more so than abundance. These results indicate that geospatial techniques and molecular gut content analysis can be combined to determine the role of land use in influencing predator–prey interactions and resulting predation events in agroecosystems.     

Relative growth rates of predator and prey dinosaurs reflect effects of predation

Hadrosaurs grew rapidly, and quantifying their growth is key to understanding life-history interactions between predators and prey during the Late Cretaceous. In this study, we longitudinally sampled a sequence of lines of arrested growth (LAGs) from an essentially full-grown hadrosaur Hypacrosaurus stebingeri (MOR 549). Spatial locations of LAGs in the femoral and tibial transverse sections of MOR 549 were measured and circumferences were calculated. For each bone, a time series of circumference data was fitted to several stochastic, discrete growth models. Our results suggest that the femur and the tibia of this specimen of Hypacrosaurus probably followed a Gompertz curve and that LAGs reportedly missing from early ontogeny were obscured by perimedullary resorption. In this specimen, death occurred at 13 years and took approximately 10-12 years to reach 95 per cent asymptotic size. The age at growth inflection, which is a proxy for reproductive maturity, occurred at approximately 2-3 years. Comparisons with several small and large predatory theropods reveal that MOR 549 grew faster and matured sooner than they did. These results suggest that Hypacrosaurus was able to partly avoid predators by outgrowing them.     

Density-dependent prey behaviours and mutable predator foraging modes induce Allee effects and over-prediction of prey mortality rates

1. Predator–prey models are often used to represent consumptive interactions between species but, typically, are derived using simple experimental systems with little plasticity in prey or predator behaviours. However, many prey and predators exhibit a broad suite of behaviours. Here, we experimentally tested the effect of density-dependent prey and predator behaviours on per capita relative mortality rates using Florida bass (Micropterus floridanus) consuming juvenile Bluegill (Lepomis macrochirus).
2. Experimental ponds were stocked with a factorial design of low, medium, and high prey and predator densities. Prey mortality, prey–predator behaviours, and predator stomach contents were recorded over or after 7 days. We assumed the mortality dynamics followed foraging arena theory. This pathologically flexible predator–prey model separates prey into invulnerable and vulnerable pools where predators can consume prey in the latter. As this approach can represent classic Lotka–Volterra and ratio-dependent dynamics, we fit a foraging arena predator–prey model to the number of surviving prey.
3. We found that prey exhibited density-dependent prey behaviours, hiding at low densities, shoaling at medium densities, and using a provided refuge at high densities. Predators exhibited ratio-dependent behaviours, using an ambush foraging mode when one predator was present, hiding in the shadows at low prey–high predator densities, and shoaling at medium and high prey–high predator densities. The foraging arena model predicted the mortality rates well until the high prey–high predator treatment where group vigilance prey behaviours occurred and predators probably interfered with one another resulting in the model predicting higher mortality than observed.
4. This is concerning given the ubiquity of predator–prey models in ecology and natural resource management. Furthermore, as Allee effects engender instability in population regulation, it could lead to inaccurate predictions of conservation status, population rebuilding or harvest rates.

     

Dynamics of predator and modular prey: effects of module consumption on stability of prey–predator system

In traditional models of predator–prey population dynamics, it is usually assumed that consumed prey are immediately removed from the population. However, in plant–herbivore interactions, damaged plants are generally alive after attacks by herbivores. This can result in successive or simultaneous attacks by multiple predators on a single prey item (here, the term prey is expanded to include plants). We constructed a mathematical model with two time scales, taking into account predation processes within a generation, considering post‐predation survival and the modularity of prey. We assumed that a prey item can be divided into modules and that it can be fed on by multiple predators or parasitized by multiple parasites at the same time. The model includes two essential factors: the modularity of prey for predators (n) and the detaching/attaching ratio of predators to prey (ε). Based on the formulae, we revealed a general property of realistic dynamics in plant–herbivore and host–parasite interactions. The analysis showed that the model could be approximated by models with the type I, type II or Beddington–DeAngelis functional responses by taking appropriate limits to the situations. When modularity is low or the detaching/attaching ratio is high, population dynamics tend to be stabilized. These stabilizing effects may be related to interference competition among predator individuals or increases in free prey modules and free predator individuals. In the limit of high modularity, the ratio of the attached prey modules to the total prey modules becomes negligible and the dynamics tend to be destabilized. However, if quantity and quality of prey modules are negatively correlated, the equilibrium is likely to be stabilized at high modularity as long as it remains feasible. These results suggest that considering post‐predation survival and modularity of prey is crucial to understand predator–prey interactions.     

Density-dependent effects of prey defenses and predator offenses

Defenses protect prey, while offenses arm predators. Some defenses and offenses are constitutive (e.g. tortoise shells), while others are phenotypically plastic and not always expressed (e.g. neckteeth in water fleas). All of them are costly and only adaptive at certain prey densities. Here, I analyse such density-dependent effects, applying a functional response model to categorize defenses and offenses and qualitatively predict at which prey densities each category should evolve (if it is constitutive) or be expressed (if it is phenotypically plastic). The categories refer to the step of the predation cycle that a defense or offense affects: (1) search, (2) encounter, (3) detection, (4) attack, or (5) meal. For example, prey warning signals such as red coloration prevent predator attacks and are hence step 4 defenses, while sharp predator eyes enhance detection and are step 3 offenses. My theoretical analyses predict that step 1 defenses, which prevent predators from searching for their next meal (e.g. toxic substances), evolve or are expressed at intermediate prey densities. Other defenses, however, should be most beneficial at low prey densities. Regarding predators, step 1 offenses (e.g. immunity against prey toxins) are predicted to evolve or be expressed at high prey densities, other offenses at intermediate densities. I provide evidence from the literature that supports these predictions.     

Selectivity underlies the dissociation between seasonal prey availability and prey consumption in a generalist predator

Generalist predators are capable of selective foraging, but are predicted to feed in close proportion to prey availability to maximize energetic intake especially when overall prey availability is low. By extension, they are also expected to feed in a more frequency‐dependent manner during winter compared to the more favourable foraging conditions during spring, summer and fall seasons. For 18 months, we observed the foraging patterns of forest‐dwelling wolf spiders from the genus Schizocosa (Araneae: Lycosidae) using PCR‐based gut‐content analysis and simultaneously monitored the activity densities of two common prey: springtails (Collembola) and flies (Diptera). Rates of prey detection within spider guts relative to rates of prey collected in traps were estimated using Roualdes’ c_st model and compared using various linear contrasts to make inferences pertaining to seasonal prey selectivity. Results indicated spiders foraged selectively over the course of the study, contrary to predictions derived from optimal foraging theory. Even during winter, with overall low prey densities, the relative rates of predation compared to available prey differed significantly over time and by prey group. Moreover, these spiders appeared to diversify their diets; the least abundant prey group was consistently overrepresented in the diet within a given season. We suggest that foraging in generalist predators is not necessarily restricted to frequency dependency during winter. In fact, foraging motives other than energy maximization, such as a more nutrient‐focused strategy, may also be optimal for generalist predators during prey‐scarce winters.     

Scale dependent effects of native prey diversity,prey biomass and natural disturbance on the invasion success of an exotic predator

There is growing support for the general notion that the drivers of invasion success often shift from biotic to abiotic factors with increasing spatial scale. Most of this research, however, has been conducted on a single trophic level; i.e. it has primarily looked at how the diversity of native competitors may influence invasion success. Less attention has been paid to understanding how native prey diversity may influence the invasion success of exotic predators and whether such biotic factors are scale-dependent. We used a hierarchical spatial survey of 17 stream communities to test whether native prey diversity, along with native prey biomass, algal resource abundance and annual stream discharge, influenced the abundance of an exotic crayfish predator, and whether the importance of these factors were scale-dependent. We used a hierarchical generalized linear model to evaluate the influence of these community and stream characteristics on exotic crayfish abundance at both the transect scale (1 m²) and the stream scale (400 m²). Our results indicated that at the stream scale, high stream discharge significantly limited invader abundance. However, at the smaller transect scale, native prey biomass was a significant driver of invasion success and positively correlated with invader abundance. We suggest that our results add to the emerging pattern that abiotic processes are stronger determinants of invasion success at large spatial scales, whereas biotic processes become more important with decreasing spatial scale. However, for predator invasions, prey biomass, not prey diversity may be a more important for driver of invasion success at small spatial scales.     

Comparison of native and non‐native predator consumption rates and prey avoidance behavior in North America and Europe

Novel predator–prey interactions can contribute to the invasion success of non‐native predators. For example, native prey can fail to recognize and avoid non‐native predators due to a lack of co‐evolutionary history and cue dissimilarity with native predators. This might result in a competitive advantage for non‐native predators. Numerous lady beetle species were globally redistributed as biological control agents against aphids, resulting in novel predator–prey interactions. Here, we investigated the strength of avoidance behavior of the pea aphid (Acyrthosiphon pisum) toward chemical cues of native lady beetles and non‐native Asian Harmonia axyridis and European Coccinella septempunctata and Hippodamia variegata in North America, hypothesizing that cues of non‐native lady beetles induce weaker avoidance behavior than cues of co‐evolved native lady beetles. Additionally, we compared aphid consumption of lady beetles, examining potential predation advantages of non‐native lady beetles. Finally, we compared cue avoidance behavior between North American and European pea aphid populations and aphid consumption of native and non‐native lady beetles in North America and Europe. In North America, pea aphids avoided chemical cues of all ladybeetle species tested, regardless of their origin. In contrast to pea aphids in North America, European pea aphids did not avoid cues of the non‐native H. axyridis. The non‐native H. axyridis and C. septempunctata were among the largest and most voracious lady beetle species tested, on both continents. Consequently, in North America non‐native lady beetle species might have a competitive advantage on shared food resources due to their relatively large body size, compared to several native American lady beetle species. In Europe, however, non‐native H. axyridis might benefit from missing aphid cue avoidance as well as a large body size. The co‐evolutionary time gap between the European and North American invasion of H. axyridis likely explains the intercontinental differences in cue avoidance behavior and might indicate evolution in aphids toward non‐native predators.     

Functional benefits of predator species diversity depend on prey identity

Abstract.  1. Determining the functional significance of species diversity in natural enemy assemblages is a key step towards prediction of the likely impact of biodiversity loss on natural pest control processes. While the biological control literature contains examples in which increased natural enemy diversity hinders pest control, other studies have highlighted mechanisms where pest suppression is promoted by increased enemy diversity.
2. This study aimed to test whether increased predator species diversity results in higher rates of predation on two key, but contrasting, insect pest species commonly found in the rice ecosystems of south-east Asia.
3. Glasshouse experiments were undertaken in which four life stages of a planthopper ( Nilaparvata lugens ) and a moth ( Marasmia patnalis ) were caged with single or three-species combinations of generalist predators.
4. Generally, predation rates of the three-species assemblages exceeded expectation when attacking M. patnalis , but not when attacking N. lugens. In addition, a positive effect of increased predator species richness on overall predation rate was found with M. patnalis but not with N. lugens .
5. The results are consistent with theoretical predictions that morphological and behavioural differentiation among prey life stages promotes functional complementarity among predator species. This indicates that emergent species diversity effects in natural enemy assemblages are context dependent; they depend not only on the characteristics of the predators species, but on the identity of the species on which they prey.     

The impact of a sit-and-wait predator: separating consumption and prey emigration

Reviews of the impact of invertebrate predators in enclosure/exclosure experiments suggest that much of the apparent depletion of prey is due to prey emigration induced by the predators. However, these generalisations derive mainly from studies of invertebrate predators that are predominantly active searchers (usually stoneflies) and of prey with strong avoidance responses (mainly mayflies).
We examined the impact of a large sit-and-wait predator, the nymph of the dragonfly Cordulegaster boltonii , which has recently invaded Broadstone Stream as a new top predator. Field enclosure/exclosure experiments were conducted to assess the impact of the invader on the benthos. Depletion of prey varied seasonally and among taxa, and was highest when prey density and encounter rates were high. Mobile prey, although least likely to show a statistically significant response because of high exchange rates, were those most strongly depleted.
Experimental channels were used to separate the relative contribution of consumption and emigration to total impact for the two most depleted prey species. Depletion of prey was due solely to consumption and predators did not induce emigration. We therefore urge caution in making generalisations about the impacts of invertebrate predators, since sit-and-wait and searching predators potentially have very different impacts.     

No evidence of nonlinear effects of predator density,refuge availability,or body size of prey on prey mortality rates

Predator density, refuge availability, and body size of prey can all affect the mortality rate of prey. We assume that more predators will lead to an increase in prey mortality rate, but behavioral interactions between predators and prey, and availability of refuge, may lead to nonlinear effects of increased number of predators on prey mortality rates. We tested for nonlinear effects in prey mortality rates in a mesocosm experiment with different size classes of western mosquitofish (Gambusia affinis) as the prey, different numbers of green sunfish (Lepomis cyanellus) as the predators, and different levels of refuge. Predator number and size class of prey, but not refuge availability, had significant effects on the mortality rate of prey. Change in mortality rate of prey was linear and equal across the range of predator numbers. Each new predator increased the mortality rate by about 10% overall, and mortality rates were higher for smaller size classes. Predator–prey interactions at the individual level may not scale up to create nonlinearity in prey mortality rates with increasing predator density at the population level.     

Detection and decay rates of prey and prey symbionts in the gut of a predator through metagenomics

DNA methods are useful to identify ingested prey items from the gut of predators, but reliable detection is hampered by low amounts of degraded DNA. PCR‐based methods can retrieve minute amounts of starting material but suffer from amplification biases and cross‐reactions with the predator and related species genomes. Here, we use PCR‐free direct shotgun sequencing of total DNA isolated from the gut of the harlequin ladybird Harmonia axyridis at five time points after feeding on a single pea aphid Acyrthosiphon pisum. Sequence reads were matched to three reference databases: Insecta mitogenomes of 587 species, including H. axyridis sequenced here; A. pisum nuclear genome scaffolds; and scaffolds and complete genomes of 13 potential bacterial symbionts. Immediately after feeding, multicopy mtDNA of A. pisum was detected in tens of reads, while hundreds of matches to nuclear scaffolds were detected. Aphid nuclear DNA and mtDNA decayed at similar rates (0.281 and 0.11 h^?1 respectively), and the detectability periods were 32.7 and 23.1 h. Metagenomic sequencing also revealed thousands of reads of the obligate Buchnera aphidicola and facultative Regiella insecticola aphid symbionts, which showed exponential decay rates significantly faster than aphid DNA (0.694 and 0.80 h^?1, respectively). However, the facultative aphid symbionts Hamiltonella defensa, Arsenophonus spp. and Serratia symbiotica showed an unexpected temporary increase in population size by 1–2 orders of magnitude in the predator guts before declining. Metagenomics is a powerful tool that can reveal complex relationships and the dynamics of interactions among predators, prey and their symbionts.     

Environmental change and predator diversity drive alpha and beta diversity in freshwater macro and microorganisms

Global biodiversity is eroding due to anthropogenic causes, such as climate change, habitat loss, and trophic simplification of biological communities. Most studies address only isolated causes within a single group of organisms; however, biological groups of different trophic levels may respond in particular ways to different environmental impacts. Our study used natural microcosms to investigate the predicted individual and interactive effects of warming, changes in top predator diversity, and habitat size on the alpha and beta diversity of macrofauna, microfauna, and bacteria. Alpha diversity (i.e., richness within each bromeliad) generally explained a larger proportion of the gamma diversity (partitioned in alpha and beta diversity). Overall, dissimilarity between communities occurred due to species turnover and not species loss (nestedness). Nevertheless, the three biological groups responded differently to each environmental stressor. Microfauna were the most sensitive group, with alpha and beta diversity being affected by environmental changes (warming and habitat size) and trophic structure (diversity of top predators). Macrofauna alpha and beta diversity was sensitive to changes in predator diversity and habitat size, but not warming. In contrast, the bacterial community was not influenced by the treatments. The community of each biological group was not mutually concordant with the environmental and trophic changes. Our results demonstrate that distinct anthropogenic impacts differentially affect the components of macro and microorganism diversity through direct and indirect effects (i.e., bottom‐up and top‐down effects). Therefore, a multitrophic and multispecies approach is necessary to assess the effects of different anthropogenic impacts on biodiversity.     

Intraguild interactions among generalist predator functional groups drive impact on herbivore and decomposer prey

Different functional groups of generalist predators may complement each other in controlling prey populations; but intraguild interactions, common among generalist predators, may also reduce the strength of top–down control. In natural communities greater alterations to ecosystem function are expected if a whole functional group declines in abundance or is lost. Therefore studying functional group diversity is important for predicting effects of predator loss. We studied the top–down impact of web‐building spiders, hunting spiders and ants, which are highly abundant generalist predators in most terrestrial ecosystems, on prey from the herbivore and decomposer system of a grassland food web. The density of the three predator groups was manipulated by continuous removal in a three‐factorial designed field experiment, which was carried out for two years. We found no positive effect of increasing predator functional group richness on prey control. However there was evidence for strong composition effects between the functional groups. The presence of ants in predator assemblages reduced the prey suppression through mostly trait‐mediated intraguild interactions, while hunting and web‐building spiders contributed additively to prey suppression and reduced the density of herbivore and decomposer prey by 50–60%. A trophic cascade on plant biomass triggered by web‐builders and hunting spiders was diminished at levels of higher predator group diversity. In conclusion, our experiments showed that intraguild interactions strongly influence the strength of top–down control by generalist predators. Among spiders there was evidence for a positive relation between functional group richness and prey suppression but the overall outcome strongly depended on the occurrence of interference, driven by trait‐mediated indirect interactions.     

Multiple predator effects result in risk reduction for prey across multiple prey densities

Investigating how prey density influences a prey’s combined predation risk from multiple predator species is critical for understanding the widespread importance of multiple predator effects. We conducted experiments that crossed six treatments consisting of zero, one, or two predator species (hellgrammites, greenside darters, and creek chubs) with three treatments in which we varied the density of mayfly prey. None of the multiple predator effects in our system were independent, and instead, the presence of multiple predator species resulted in risk reduction for the prey across both multiple predator combinations and all three levels of prey density. Risk reduction is likely to have population-level consequences for the prey, resulting in larger prey populations than would be predicted if the effects of multiple predator species were independent. For one of the two multiple predator combinations, the magnitude of risk reduction marginally increased with prey density. As a result, models predicting the combined risk from multiple predator species in this system will sometimes need to account for prey density as a factor influencing per-capita prey death rates.     

Effects of plant gross morphology on predator consumption rates

We find that spatial structure, and in particular, differences in gross plant morphology, can alter the consumption rates of generalist insect predators. We compared Asian lady beetle, Harmonia axyridis Pallas, and green lacewing larvae, Chrysoperla carnea Stephens, consumption rates of pea aphids, Acyrthosiphon pisum Harris, in homogeneous environments (petri dishes) and heterogeneous environments (whole plants). Spatial complexity is often described as reducing predator success, and we did find that predators consumed significantly more aphids on leaf tissue in petri dishes than on whole plants with the same surface area. However, subtle differences in plant morphology may have more unexpected effects. A comparison of consumption rates on four different isogenic pea morphs (Pisum sativum L.) controlled for surface area indicated that both lady beetles and lacewings were more successful on morphologies that were highly branched. We speculate that predators move more easily over highly branched plants because there are more edges to grasp.     

Consumer effects decline with prey diversity

While consumer species diversity is known to influence the capture of limited resources, little is known about how prey diversity impacts the transfer of energy and matter among trophic levels. Here, we perform a meta‐analysis of experiments that have examined the impact of grazers on the biomass of periphytic algae to test the hypothesis that the magnitude of consumer (grazer) effects on prey (algae) depends on the species diversity of the prey assemblage. The analysis reveals that consumer effects tend to decrease as the diversity of a prey assemblage increases. This trend is robust for several different, yet complementary indices of grazer effect size and algal diversity. The trend also remains significant after statistically controlling for a variety of factors that can covary with prey diversity among studies. We discuss several possible mechanisms for the documented pattern, such as diversity enhancing the probability of inedibility and of positive interactions.     

Evolutionary rates do not drive latitudinal diversity gradients

Among the several hypotheses invoked for explaining latitudinal diversity gradients (LDGs), some models classified within the 'evolutionary hypothesis' family assume that LDGs are the direct consequence of latitudinal variations in the speciation and/or extinctions rates. Spatially structured simulations of the biogeographical dispersal of a randomly generated clade refute the central tenet of these explanatory models and indicate that global diversity patterns are combined outcomes of geographic and thermal mid-domain effects under a phylogenetically controlled niche conservatism constraint. The positive correlation observed in several higher taxa between speciation rate and diversity does not involve any causal relationship between these two parameters but is most likely the first order by-product of a positive correlation between temperature and per capita speciation rate.