Propensity towards cannibalism among Hypoaspis aculeifer and H. miles, two soil-dwelling predatory mite species

In biological control programmes, the two predatory soil mites Hypoaspis aculeifer and H. miles are often applied against soil-borne pests like mushroom flies, springtails and mites. Although the mites show high consumption rates on varying prey types in Petri dish experiments and in greenhouses, their overall efficiency is sometimes limited. We hypothesized that intraspecific interactions, like cannibalism, could contribute to this decreased competence. Therefore, experiments were conducted to show the propensity of H. aculeifer and H. miles to cannibalise. Adult mites and nymphs were introduced as predators with conspecific eggs, larvae, nymphs, adult females or males as prey and the number of killed individuals was recorded. Additionally, the oviposition rate on conspecific prey was quantified and the correlation with the number of prey consumed was calculated to assess the influence of cannibalism on egg production. The results illustrate that cannibalism occurs infrequently in both Hypoaspis spp., the only exception being H. aculeifer nymphs, which cannibalised one conspecific egg per day. Moreover, cannibalism never occurred in the presence of alternative prey. Oviposition rate decreased during the experiment in both species but it was positively correlated with the cannibalism rate only for H. aculeifer. The benefit of cannibalism for populations of H. aculeifer and H. miles is discussed.     

Diet, morphology, and interspecific killing in carnivora

Interspecific killing is a key determinant of the abundances and distributions of carnivores, their prey, and nonprey community members. Similarity of body size has been proposed to lead competitors to seek similar prey, which increases the likelihood of interference encounters, including lethal ones. We explored the influence of body size, diet, predatory habits, and taxonomic relatedness on interspecific killing. The frequency of attacks depends on differences in body size: at small and large differences, attacks are less likely to occur; at intermediate differences, killing interactions are frequent and related to diet overlap. Further, the importance of interspecific killing as a mortality factor in the victim population increases with an increase in body size differences between killers and victims. Carnivores highly adapted to kill vertebrate prey are more prone to killing interactions, usually with animals of similar predatory habits. Family-level taxonomy influences killing interactions; carnivores tend to interact more with species in the same family than with species in different families. We conclude that although resource exploitation (diet), predatory habits, and taxonomy are influential in predisposing carnivores to attack each other, relative body size of the participants is overwhelmingly important. We discuss the implications of interspecific killing for body size and the dynamics of geographic ranges.     

Biological Control of the Bulb Mite, Rhizoglyphus robini , by the Predatory Mite, Hypoaspis aculeifer , on Lilies: Predator-Prey Dynamics in the Soil, under Greenhouse and Field Conditions

We tested the capacity of the soil-dwelling predatory mite, Hypoapsis aculeifer , to control mites attacking lily bulbs. Experiments in the greenhouse and in the field showed that in the absence of predatory mites populations of the bulb mite, Rhizoglyphus robini , on lily bulbs increased, whereas the release of predatory mites either slowed down the increase - as observed in the field - or caused the bulb mites populations to decrease - as observed in the greenhouse. In all cases the population of predatory mites increased as long as bulb mite densities were not too low. However, within the first week after predator release there was usually a sharp decline to 10-40% of the original number released. Greenhouse experiments on intact lily bulbs in pots, boxes and 1 m 2 plots with peat soil showed that when released in a ratio of 1 predator to 2 or 5 prey, the predatory mite, Hypoaspis aculeifer , suppressed populations of bulb mites to less than 10 individual per bulb within 6 weeks. Elimination of bulb mites was observed only when the predator-to-prey ratio at release was equal to 3:1. Field experiments in 2 m 2 plots with intact bulbs in rather compact sandy soil showed that when released in ratio of 1 predator to 1 or 2 prey, the predatory mite, H. aculeifer , did not cause the population of bulb mites to decrease, but it did reduce their population growth. The initial predator-to-prey ratios required to achieve suppression (ca 1:2) or elimination (3:1) in the soil environment are much higher than those required for bulb mite elimination when lily bulb scales were embedded in a medium of vermiculite (ca 1:20). Among the possible causes are: (1) the initial losses of predators in the greenhouse and even more so in the field due to mortality and/or emigration from the experimental plots; (2) the lower temperatures in the greenhouse and especially in the field, which slow down the growth and predation processes and thereby delay prey extinction; and (3) the spatial complexity of the soil environment which creates refuges for the bulb mites.     

Effects of satiation and starvation on nematocyst discharge, prey killing, and ingestion in two species of sea anemone

Studies spanning 60 years with several cnidarian species show that satiation inhibits prey capture and ingestion and that starvation increases prey capture and ingestion. Most have attributed the effects of satiation to inhibition of nematocyst discharge. We hypothesized that satiation inhibits prey capture and ingestion in sea anemones (Haliplanella luciae and Aiptasia pallida) primarily by inhibiting the intrinsic adherence (i.e., holding power) of discharging nematocysts. Using a quantitative feeding assay for H. luciae, we found that satiation completely uncoupled prey killing from prey ingestion, while nematocyst-mediated prey killing was only partially inhibited. Using A. pallida to measure nematocyst discharge and nematocyst-mediated adhesive force, we showed that satiation completely inhibited the intrinsic adherence of discharging nematocysts from Type B and Type C cnidocyte/supporting cell complexes (CSCCs), while only partially inhibiting nematocyst discharge from Type Bs. These inhibitory effects of satiation were gradually restored by starvation, reaching a maximum at 72 h after feeding. Thus, the effects of satiation and starvation on prey killing and ingestion in two species of acontiate sea anemones are primarily due to changes in the intrinsic adherence of nematocysts from both Type B and Type C CSCCs.     

The biology of Pholcus phalangioides (Araneae, Pholcidae): predatory versatility, araneophagy and aggressive mimicry

Predatory versatility occurs in Pholcus phalangioides (Fuesslin). In addition to building prey-catching space webs, P. phalangioides invades webs of other spiders and feeds on the occupants. It acts as an aggressive mimic by performing specialized vibratory behaviours to which the prey-spider responds as it normally would to its own prey. Prey (spiders and insects) is attacked by wrapping. Prey that trips over lines at the edge of a web of P. phalangioides , but fails to enter the web, is successfully attacked: P. phalangioides leans out of its web to throw silk over the prey, keeping as few as two legs on the silk. However, P. phalangioides does not attack prey that is completely away from webs. Occasionally, P. phalangioides feeds on eggs of other spiders and on ensnared insects it encounters in alien webs. Experimental evidence indicates that vision is of little or no importance in the predatory behaviour of P. phalangioides . Although P. phalangioides invades diverse types of webs, in addition to using its own web, its efficiency as a predator varies with web-type. It is most efficient as a predator of spiders and, especially, insects on its own web, and least efficient as a predator of amaurobiids on their cribellate sheet webs. Sensory, locomotory and other factors which influence differential predatory efficiency are discussed. The behaviour of P. phalangioides is compared to that of Portia , an araneophagic web-invading salticid, and the results of this study are discussed in relation to hypotheses concerning salticid evolution.     

Combining plant- and soil-dwelling predatory mites to optimise biological control of thrips

The efficiency of a natural enemy combination compared to a single species release for the control of western flower thrips (WFT) Frankliniella occidentalis (Pergande) on cucumber plants was investigated. Since a large part of F occidentalis seems to enter the soil passage, a joint release of the plant-inhabiting predatory mite Amblyseius cucumeris (Oudemans) that feeds on thrips first-instar larvae and the soil-dwelling predatory mite Hypoaspis aculeifer (Canestrini) that preys on thrips pupae in the ground might offer a promising approach for a holistic control strategy. Therefore, two sets of experiments were conducted in cooperation with a commercial vegetable grower where the plants in plots were infested with a defined number of larval and adult F occidentalis. Two species of natural enemies were released either synchronously or solely, and their efficacy was compared to control plots devoid of antagonists. In both experiments, the predatory mites were released twice with a density of 46 A. cucumeris/m2, and 207 H. aculeifer/m2 (low-density) in the first experiment and 528 H. aculeifer/m2 (high-density) in the second one. Population growth of all arthropod species on the plants and in the soil was quantified at regular intervals and included all soil-dwelling mites and alternative preys present in the substrate. The results showed that H. aculeifer alone had a significant impact on thrips population development only when released at high-densities, but competence was lower compared to the other antagonist treatments. The impact of A. cucumeris alone and A. cucumeris & H. aculeifer combined was similar. Thus, the pooled exploitation of natural enemies did not boost thrips control compared to the single species application of A. cucumeris (non-additive effect), which could be explained by resource competition between both predatory mite species. Species number and population size in the soil of the experimental plots both showed a high variability, a possible consequence of their interaction with released soil-dwelling predatory H. aculeifer mites. The impact of resource competition and presence of alternative preys on thrips biological control is exhaustively discussed. From our study, we can extract the subsequent conclusions: (1) the combined use of H. aculeifer and A. cucumeris cannot increase thrips control on cucumber compared to the release of A. cucumeris alone, but the overall reliability of thrips biological control might be enhanced, (2) the availability of alternative preys seemed to affect the thrips predation rate of H. aculeifer, and (3) the impact of naturally occurring soil predatory mites on the control of WFT seemed to be partial.     

The influence of turbulence on plankton predation strategies

The importance of predation in regulating the size of competing plankton and larval fish populations has long been appreciated. However, it has only recently been recognized that turbulence must have a significant influence on predator-prey interactions because most rival species of microorganisms co-exist in oceanic or fast moving fresh water flows. Turbulence is likely to influence predation strategies in two ways. The extra energy imparted to a micro-organism from the flow field will enhance the number of encounters or "contacts" between predators and prey. At the same time, because the velocity of a predator relative to its potential prey will be increased, the time-scale over which a capture must be completed is reduced. Balancing the benefits of extra encounters with the drawbacks of more difficult captures, will dictate an optimal predation strategy, either foraging behaviour or ambush feeding, on the predator. This will depend on its own and the prey's swimming capabilities, as well as the characteristics of the turbulent environment. In this paper some previous work, examining the increased encounter rate in turbulence, will be extended to look at the capture problem. The main proposal is that the capture event should be encapsulated in a capture probability function, from which the optimal predation strategy can be derived. As an illustration, plausible capture probability functions will be postulated and the resulting predictions tested against numerical simulations carried out in a turbulent-like flow field. Good agreement between the predictions and the simulations is demonstrated.     

The effects of overwintering and habitat type on body condition and locomotion of the wolf spider Pardosa alacris

Overwintering in temperate regions is a prominent mortality risk for invertebrates and may affect their behaviour and body condition. Pardosa alacris is a common ground dwelling spider in central European native and plantation forests, and habitat type and prey availability may play important roles in their overwintering. The effect of overwintering on body condition and behaviour of spiders in semi natural and exotic habitats is relatively unknown. Here we assess the effects of winter on spiders from native poplar and exotic pine plantations. The locomotory behaviour of P. alacris (distance covered and speed) was assessed by tracking their movement in a white circular plastic arena. We assessed body condition, body size, and total fat content. Forest type and sex had significant effects on body length. Fat content was significantly higher in the spring than in autumn, and spiders covered larger distances and were faster in autumn than in spring. Fat content had a significant negative effect on average speed. Spiders in native forests were smaller but grew more during the winter than in exotic plantations, possibly due to higher prey availability in native forests. Visually-hunting predators may significantly affect spiders. Fat spiders with better body condition moved less, and were thus less detectable by predators. However the low movement rate may result in a low rate of encountering prey items, thus lowering feeding efficiency.     

Locking out predators by silk,a new counterattack behaviour in a social spider mite

1. A type of arms race that includes predation, counterattacks and cross‐counterattacks occurs between the phytophagous mite Stigmaeopsis nanjingensis (Ma et Yuan), which lives in self‐woven nests and exhibits cooperative sociality, and its specialised phytoseiid mite predator, Typhlodromus bambusae Ehara. 2. First, the efficiency of the S. nanjingensis (prey) counterattacking T. bambusae (predator) was observed. The prey females frequently locked the immature predators out of their nests using silk web, and the predators subsequently died of starvation. Furthermore, the prey males often killed immature T. bambusae mites after they invaded the nests. 3. This reversal of roles in the predator–prey system was then re‐reversed (returned to a normal state) by the behaviour of T. bambusae females. Immature predators could maintain their predacious natures due to the presence of attending adult females, which are able to cope with the prey counterattack behaviours.     

The foraging ecology of larval and juvenile fishes

Knowledge of the foraging ecology of fishes is fundamental both to understanding the processes that function at the individual, population and community levels, and for the management and conservation of their populations and habitats. Furthermore, the factors that influence the acquisition and assimilation of food can have significant consequences for the condition, growth, survival and recruitment of fishes. The majority of marine and freshwater fish species are planktivorous at the onset of exogenous nutrition and have a limited ability to detect, capture, ingest and digest prey. Improvements in vision, development of fins and associated improvements in swimming performance, increases in gape size and development of the alimentary tract during ontogeny often lead to shifts in diet composition. Prey size, morphology, behaviour and abundance can all influence the prey selection of larval and juvenile fishes. Differences in feeding behaviour between fish species, individuals or during ontogeny can also be important, as can inter- and intraspecific interactions (competition, predation risk). Temporal (diel, seasonal, annual) and spatial (microhabitat, mesohabitat, macrohabitat, regional) variations in prey availability can have important implications for the prey selection, diet composition, growth, survival, condition and, ultimately, recruitment success of fishes. For fish populations to persist, habitat must be available in sufficient quality and quantity for the range of activities undertaken during all periods of development. Habitats that enhance the diversity, size ranges and abundance of zooplankton should ensure that sufficient food resources are available to larval and juvenile fishes.     

Effect of starvation time on the prey capture behaviour, functional response and population growth of Asplanchna sieboldi (Rotifera)

1. We examined the effect of different periods of prior starvation(from 30 min to 16 h) on the prey capture behaviour, and functional and numerical responses of the predatory rotifer Asplanchna sieboldi using the rotifer Brachionus calyciflorus as prey.
2. Feeding activity (i.e. encounter, attack, capture and ingestion) by Asplanchna increased significantly with increasing prey densities from 2 to 16 mL⁻⁻¹ and with increasing prior starvation periods from 0.5 to 8 h.
3.  Asplanchna sieboldi showed a type II functional response at all the prior starvation periods tested. The asymptotic prey density was highest after 8 h of starvation.
4. The instantaneous population growth rate of A. sieboldi ranged from 0.089 ± 0.044 (when starved for 8 h in every 24 h and at a prey density of 2 individuals mL⁻⁻¹ for the other 16 h) to 1.015 ± 0.142 in the control (no starvation and at a prey density of 16 individuals mL⁻⁻¹). The effect of starvation time on the numerical response was evident only at the higher prey density.     

Prey concealment and spatiotemporal patrolling behaviour of the Indian tree ant Oecophylla smaragdina (Fabricius)

Summary: Oecophylla smaragdina workers conceal prey with leaves and twigs on discovering large prey on their ground territory. The prey concealment behaviour occurs concurrently with prey capture and killing. Ants were observed conducting spatiotemporal patrolling around large prey. The garden lizard Calotes versicolor and at least two predatory ant species were found to forage in the same area as O. smaragdina. It is suggested that the prey concealment behaviour of Oecophylla may be a strategy to prevent prey detection by vertebrate predators which use vision to prey on similar large prey species.     

Multi-behavioral strategies in a predator–prey game: an evolutionary algorithm analysis

Behavioral games between predators and prey often involve two sub-games: 'pre-encounter' games affecting the rate of encounter between predators and prey (e.g. predator–prey space games, Sih 2005 ), and 'post-encounter' games that influence the outcome of encounters (e.g. waiting games at prey refugia, Hugie 2003 , and games of vigilance, Brown et al. 1999 ). Most models, however, focus on only one or the other of these two sub-games.
I investigated a multi-behavioral game between predators and prey that integrated both pre-encounter and post-encounter behaviors. These behaviors included landscape-scale movements by predators and prey, a type of prey vigilance that increases immediately after an encounter and then decays over time ('ratcheting vigilance'), and predator management of prey vigilance. I analyzed the game using a computer-based evolutionary algorithm. This algorithm embedded an individual-based model of ecological interactions within a dynamic adaptive process of mutation and selection. I investigated how evolutionarily stable strategies (ESS) varied with the predators' learning ability, killing efficiency, density and rate of movement. I found that when predators learn prey location, random prey movement can be an ESS. Increased predator killing efficiency reduced prey movement, but only if the rate of predator movement was low. Predators countered ratcheting vigilance by delaying their follow-up attacks; however, this delay was reduced in the presence of additional predators. The interdependence of pre-and post-encounter behaviors revealed by the evolutionary algorithm suggests an intricate co-evolution of multi-behavioral predator–prey behavioral strategies.     

Resource dynamics influence the strength of non-consumptive predator effects on prey

Predators influence prey populations both by consuming individual prey, and by inducing changes in prey behaviour that limit reproduction and survival. Because prey trade-off predation risk for forageing gains, the magnitude of predators' non-consumptive effects should depend on resource availability. Studies of non-consumptive effects generally adopt either of two strategies: (i) maintaining a static ration of the prey's resources; and (ii) using resource populations that vary dynamically in response to prey behaviour. Contrasting these experimental designs using meta-analysis, we evaluated whether resource dynamics influence the magnitude of non-consumptive effects on prey growth, survival, fecundity, population density, forageing rate and habitat use. Predators had a more negative effect on prey demography in dynamic- vs. static-resource experiments. Our results highlight the importance of resource dynamics in mediating the magnitude of non-consumptive effects of predators on prey, and illustrate the often-unintended impacts of experimental design on estimates of effect size in ecological interactions.     

The behaviour of 9 stored product beetles at pitfall trap arenas and their capture in millet

Pitfall trap capture forSitophilus oryzae (L.),S. zeamais (Motschulsky),S. granarius (L.),Tribolium confusum (Duval),T. castaneum (Herbst);Oryzaephilus surinamensis (L.),Cryptolestes pusillus (Schonherr),Rhyzopertha dominica (Fabr.), andProstephanus truncatus (Horn) in millet was assessed by visual and time-lapse video recordings. The behaviour of different beetle species in arenas containing millet was monitored over 24 h and the frequency of encounters with the trap rim resulting in capture recorded. The capture efficiency of four types of pitfall traps (i.e. polystyrene, polythene, glass and tin-plated steel can) with rims exposed or submerged below the millet surface level were compared. Capture was related to beetle size, locomotory rate, and beetle behaviour at the trap rim as well as trap design and placement. The lighter and smaller species were least captured. Glass jars were more effective than plastic and metal containers. Traps placed with their rims submerged below the grain surface level were more efficient than those with rims exposed. Capture rate was unrelated to trap size. The frequency of encounters with trap rims was not correlated with capture rate. Three types of avoidance behaviours at the trap rims i.e. probing, skirting and spontaneous retreat, were related to capture rate, spontaneous retreat being the most effective escape mechanism and probing least.     

Predator-induced flow disturbances alert prey,from the onset of an attack

Many prey species, from soil arthropods to fish, perceive the approach of predators, allowing them to escape just in time. Thus, prey capture is as important to predators as prey finding. We extend an existing framework for understanding the conjoint trajectories of predator and prey after encounters, by estimating the ratio of predator attack and prey danger perception distances, and apply it to wolf spiders attacking wood crickets. Disturbances to air flow upstream from running spiders, which are sensed by crickets, were assessed by computational fluid dynamics with the finite-elements method for a much simplified spider model: body size, speed and ground effect were all required to obtain a faithful representation of the aerodynamic signature of the spider, with the legs making only a minor contribution. The relationship between attack speed and the maximal distance at which the cricket can perceive the danger is parabolic; it splits the space defined by these two variables into regions differing in their values for this ratio. For this biological interaction, the ratio is no greater than one, implying immediate perception of the danger, from the onset of attack. Particular attention should be paid to the ecomechanical aspects of interactions with such small ratio, because of the high degree of bidirectional coupling of the behaviour of the two protagonists. This conclusion applies to several other predator–prey systems with sensory ecologies based on flow sensing, in air and water.     

Functional response: rigorous estimation and sensitivity to genetic variation in prey

Holling's type II functional response is a cornerstone of community ecology and coevolutionary theory. The so‐called disc equation is the most widely used model of the type II response, yet thus far no robust experimental assessment has been achieved in any single system. Fundamental issues that remain to be assessed include whether the assumptions of the disc equation are fulfilled, whether the disc equation yields accurate estimates of predation‐related individual traits, and whether differences in disc equation parameters can capture genetic variation in prey behaviour. This paper provides a rigorous approach to all of these questions. The functional response of the predatory mite Pergamasus crassipes on three genetically distinct clones of the springtail Folsomia candida was measured at six levels of prey density in controlled conditions where prey number and arena size were concomitantly manipulated. A crucial assumption of Holling's disc equation was fulfilled by maintaining a constant prey density for the entire experimental period of predation. The timing of each attack and capture, as well as the duration of the handling time, were recorded by constant observation. We contrasted three different methods to calculate functional response curves: (1) indirect estimation of the disc equation's parameters from the number of prey killed by the end of each experimental run; (2) direct estimation of the parameters via a unique protocol of constant observation; and (3) independently deriving a function based on direct measurements of encounter rate and attack success. The basic assumptions of the disk equation were globally fulfilled. Estimations of the functional response's parameters (type II) were remarkably congruent across approach (1) and (2). A single genetic effect was detected – the relationship between the encounter rate and prey density differed significantly between clones – whereas a direct comparison of functional response across clones failed to reveal genetic variation.     

Springtail postmolt vulnerability to pseudoscorpion predation: Mechanisms and implications

Arthropod prey are expected to be more vulnerable to their predators immediately following molt. The effects of springtail (Isotoma carpenteri) postmolt vulnerability on interactions with a pseudoscorpion predator were examined in the laboratory. Springtails exposed to vials pretreated with pseudoscorpions (Apochthonius minimus) delayed molting compared to those prey that were exposed to vials pretreated only with springtails. Although their escape ability (measured as distance jumped) was unaffected by molt condition, postmolt springtails were more profitable in terms of reduced predator handling time following capture. Despite this,A. minimus did not distinguish between postmolt and intermolt prey presented at either end of a T-maze.     

Prey selectivity of Piona exigua,a planktonic water mite

Summary Females, males and nymphs of Piona exigua were observed during prey capture and ingestion. The encounter radius of the mite was very small, allowing the escape of some crustaceans, such as the calanoid copepod Boeckella. Cladocerans, such as Bosmina or Chydorus, with little or no pre-contact escape response were the most vulnerable to mite predation. Preference values in size-selection experiments varied widely between individual mites. Adult mites presented with two sizes of Daphnia carinata generally preferred the smaller prey. When four sizes were presented simultaneously, however, the preferences of female mites for each size were not significantly different. Patterns of prey selection varied with predator age and sex; for example, female mites preferred Daphnia to Simocephalus, Ceriodaphnia and Chydorus, while nymphs showed a strong preference for Chydorus over Ceriodaphnia. When two prey types were present in equal proportions, differences in total prey density (range 5 or 10/1 – 30 or 50/1) did not alter preferences between the prey species. The preference of female mites for a particular prey type generally increased with increasing relative abundance of the prey type in each of three experiments (Daphnia: Ceriodaphnia, Ceriodaphnia: Chydorus, and Daphnia: Simocephalus). These results imply switching behaviour in these mites. Our results indicate the value of direct observation of predatory behaviour as an adjunct to prey selection experiments. It is also apparent that predatory behaviour in the presence of more than one prey type may not be predictable from that observed in single-prey situations. Predation rates on particular prey species were sometimes reduced in the presence of another species. The relative proportions of prey eaten when two species were present could not be predicted from the number of each species eaten when they were presented separately.     

Territorial aggressiveness and predation: two possible origins of snapping in the ant Plectroctena minor

Plectroctena minor workers have long mandibles that can snap and deliver a sharp blow to intruders or prey, stunning or killing them. Encounters between homocolonial P. minor workers separated for 24 h or 15 days never resulted in snapping, while this behaviour was always noted during encounters between heterocolonial workers on neutral arenas or on the territory of a colony. In the latter case, only the aliens, that generally tried to escape, were snapped at. Snapping also occurred during encounters with workers belonging to sympatric ponerine species. During predation, the percentages of snapping varied according to prey nature, suggesting prey discrimination. Termite soldiers were always snapped at, while other prey were more often snapped close to rather than far from the nest entrances, indicating an intermingling of territorial aggressiveness and predatory behaviour. We discuss the adaptive value of snapping for hunting in galleries.