Spatial prey availability and pulsed reproductive tactics: Encounter risk in a canid-ungulate system.

Predation risk is a function of spatiotemporal overlap between predator and prey, as well as behavioural responses during encounters. Dynamic factors (e.g. group size, prey availability and animal movement or state) affect risk, but rarely are integrated in risk assessments. Our work targets a system where predation risk is fundamentally linked to temporal patterns in prey abundance and behaviour. For neonatal ungulate prey, risk is defined within a short temporal window during which the pulse in parturition, increasing movement capacity with age and antipredation tactics have the potential to mediate risk. In our coyote-mule deer (Canis latrans-Odocoileus hemionus) system, leveraging GPS data collected from both predator and prey, we tested expectations of shared enemy and reproductive risk hypotheses. We asked two questions regarding risk: (A) How does primary and alternative prey habitat, predator and prey activity, and reproductive tactics (e.g. birth synchrony and maternal defence) influence the vulnerability of a neonate encountering a predator? (B) How do the same factors affect behaviour by predators relative to the time before and after an encounter? Despite increased selection for mule deer and intensified search behaviour by coyotes during the peak in mule deer parturition, mule deer were afforded protection from predation via predator swamping, experiencing reduced per-capita encounter risk when most neonates were born. Mule deer occupying rabbit habitat (Sylvilagus spp.; coyote's primary prey) experienced the greatest risk of encounter but the availability of rabbit habitat did not affect predator behaviour during encounters. Encounter risk increased in areas with greater availability of mule deer habitat: coyotes shifted their behaviour relative to deer habitat, and the pulse in mule deer parturition and movement of neonatal deer during encounters elicited increased speed and tortuosity by coyotes. In addition to the spatial distribution of prey, temporal patterns in prey availability and animal behavioural state were fundamental in defining risk. Our work reveals the nuanced consequences of pulsed availability on predation risk for alternative prey, whereby responses by predators to sudden resource availability, the lasting effects of diversionary prey and inherent antipredation tactics ultimately dictate risk.

Predatory mite Amblyseius orientalis prefers egg stage and low density of Carpoglyphus lactis prey.

Amblyseius orientalis (Ehara) (Acari: Phytoseiidae) is an effective predatory mite for spider mite control on fruit trees in China. In recent decades, it has been produced massively at a commercial natural enemy producer, feeding on the storage mite Carpoglyphus lactis (L.). In the predator production process, the ratio of predatory mites to their prey was found to be critical for the population increase of A. orientalis in large-scale rearings. In this study, we investigated the predatory capacity of A. orientalis on various developmental stages of the prey C. lactis, and the effect of prey numbers on predator reproduction. The maximum predation rate of A. orientalis adults on C. lactis adults was 2.21 per day at the lowest density of five prey adults, and on C. lactis eggs it was 45.07 at the highest density of 60 prey eggs. The preference index Ci of A. orientalis on C. lactis eggs and adults was 0.4312 and - 0.9249, respectively, suggesting that A. orientalis preferred eggs to adults. Amblyseius orientalis could reproduce when it preyed on either eggs or deutonymphs of C. lactis. However, the fecundity of the predatory mites is not always proportional to the provided prey number. Higher density of prey deutonymphs resulted in lower fecundity, whereas more prey eggs resulted in higher fecundity of A. orientalis. Therefore, our study indicated that the choice of suitable density and developmental stage of prey can significantly improve A. orientalis production on a large scale.

Potential of a winterschmidtiid prey mite for the production of the predatory mite Neoseiulus californicus (Acari: Phytoseiidae).

Mass rearing of the predatory mite Neoseiulus californicus (McGregor) (Acari: Phytoseiidae) using natural (prey) methods is costly and laborious, limiting its application in the biological control of pests. A high-production, low-cost method using a prey substitute would help to relieve this problem. Oulenziella bakeri Hughes (Acari: Winterschmidtiidae) could be an alternative prey source, but studies on the reproductive parameters of N. californicus under rearing conditions are lacking. This study evaluated the potential of O. bakeri as an alternative prey in N. californicus rearing by comparing developmental parameters among N. californicus reared on three diets based on an age-stage two-sex life table. We found that the preoviposition period and developmental time of N. californicus did not vary based on diet. The fecundity of N. californicus adults reared on O. bakeri was 29.8 eggs per female, which was lower than that of adults reared on Tetranychus urticae Koch (Acari: Tetranychidae) (42.9 eggs per female); there was no significant difference between O. bakeri and apple pollen (30.2 eggs per female). The oviposition rate of mites fed on O. bakeri was 69% of that fed on T. urticae. Neoseiulus californicus reared on O. bakeri and apple pollen showed the same intrinsic rate of increase (0.25 per day), which was 86% of the rate of those fed on T. urticae. Compared with predatory mites reared on natural prey, N. californicus reared on O. bakeri had a high survival rate and good oviposition and population growth parameters, suggesting that O. bakeri is suitable for the rearing of N. californicus.

Predation of boreal owl nests by pine martens in the boreal forest does not vary as predicted by the alternative prey hypothesis.

The alternative prey hypothesis (APH) states that temporally synchronous population fluctuations of microtine rodents and other small herbivores are caused by generalist predators that show functional and numerical responses to the abundance of microtines. This would lead to an increased predation of alternative prey in the low phase of the microtine population fluctuations. One candidate for such a predator is the tree-climbing pine marten (Martes martes), which includes bird eggs in its diet, among them eggs of the cavity-nesting boreal owl (Aegolius funereus). I used long-term data to test whether pine marten predation of boreal owl eggs in nest boxes varied as predicted by the APH. The probability of predation of owl nests situated < 45 km from a site where microtines were trapped in spring during four decades increased with microtine trapping index, which is opposite to the prediction from the APH. As the data set was limited to one nest per box, I extended it spatially and temporally using the clutch size of each boreal owl nest as a proxy for the actual microtine abundance at the site. The probability of nest predation increased with clutch size. However, the effects of microtine index and owl clutch size became non-significant when I controlled for habitat, and in particular cavity age, which had an overriding effect. The increase in predation probability with cavity age suggests that the long-term spatial memory of pine marten is an important factor in the pattern of its nest predation in tree cavities.

Review: predatory soil mites as biocontrol agents of above- and below-ground plant pests.

Biological pest control is becoming increasingly important for sustainable agriculture. Although many species of natural enemies are already being used commercially, efficient biological control of various pests is still lacking, and there is a need for more biocontrol agents. In this review, we focus on predatory soil mites, their role as natural enemies, and their biocontrol potential, mainly in vegetable and ornamental crops, with an emphasis on greenhouse systems. These predators are still underrepresented in biological control, but have several advantages compared to predators living on above-ground plant parts. For example, predatory soil mites are often easy and affordable to mass rear, as most of them are generalist predators, which also means that they may be used against various pests and can survive periods of pest scarcity by feeding on alternative prey or food. Many of them can also endure unfavourable conditions, making it easier for them to establish in various crops. Based on the current literature, we show that they have potential to control a variety of pests, both in greenhouses and in the field. However, more research is needed to fully understand and appreciate their potential as biocontrol agents. We review and discuss several methods to increase their efficiency, such as supplying them with alternative food and changing soil/litter structure to enable persistence of their populations. We conclude that predatory soil mites deserve more attention in future studies to increase their application in agricultural crops.

Changes in life history traits and transcriptional regulation of Coccinellini ladybirds in using alternative prey.

BACKGROUND: Ladybird beetles (Coleoptera, Coccinellidae) are highly diverse in their feeding habits. Most of them are specialist feeders, while some can have a broad spectrum of prey. As a representative group of generalists, the tribe Coccinellini includes many aphidophagous species, but members of this tribe also feed on other hemipterous insects including coccids, psyllids and whiteflies. As a result, several species are effective biological control agents or invasive species with serious non-target effects. Despite their economic importance, relatively little is known about how they adapt to new prey. RESULTS: In this study, comparisons of the life history traits and transcriptomes of ladybirds fed initial (aphids) and alternative prey (mealybugs) were performed in three Coccinellini species. The use of alternative prey greatly decreased performance, implied by the significantly prolonged development time and decreased survival rate and adult weight. Prey shifts resulted in a set of differentially expressed genes encoding chemosensory proteins and digestive and detoxifying enzymes. CONCLUSIONS: Our results suggest that these generalists do not perform well when they use alternative prey as the sole nutrition source. Although their capacity for predation might have created an opportunity to use varied prey, they must adapt to physiological obstacles including chemosensing, digestion and detoxification in response to a prey shift. These findings challenge the effect of Coccinellini predators on the biological control of non-aphid pests and suggest the possibility of non-target attacks by so-called specialists.

Non-pest prey do not disrupt aphid predation by a web-building spider.

A generalist predator's ability to contribute to biological control is influenced by the decisions it makes during foraging. Predators often use flexible foraging tactics, which allows them to pursue specific types of prey at the cost of reducing the likelihood of capturing other types of prey. When a pest insect has low nutritional quality or palatability for a predator, the predator is likely to reject that prey in favour of pursuing alternative, non-pest prey. This is often thought to limit the effectiveness of generalist predators in consuming aphids, which are of low nutritional quality for many generalist predators. Here, we report behavioural assays that test the hypothesis that the generalist predator, Grammonota inornata (Araneae: Linyphiidae), preferentially forages for a non-pest prey with high nutritional quality (springtails), and rejects a pest prey with low nutritional quality (aphids). In no-choice assays, molecular gut-content analysis revealed that spiders continued to feed on the low-quality aphids at high rates, even when high-quality springtails were readily available. When provided a choice between aphids and springtails in two-way choice tests, spiders did not show the expected preference for springtails. Decision-making by spiders during foraging therefore appears to be sub-optimal, possibly because of attraction to the less frequently encountered of two preys as part of a dietary diversification strategy. These results indicate that behavioural preferences alone do not necessarily compromise the pest-suppression capacity of natural enemies: even nutritionally sub-optimal pest prey can potentially be subject to predation and suppression by natural enemies.

Comparing rates of springtail predation by web-building spiders using Bayesian inference.

A major goal of gut-content analysis is to quantify predation rates by predators in the field, which could provide insights into the mechanisms behind ecosystem structure and function, as well as quantification of ecosystem services provided. However, percentage-positive results from molecular assays are strongly influenced by factors other than predation rate, and thus can only be reliably used to quantify predation rates under very restrictive conditions. Here, we develop two statistical approaches, one using a parametric bootstrap and the other in terms of Bayesian inference, to build upon previous techniques that use DNA decay rates to rank predators by their rate of prey consumption, by allowing a statistical assessment of confidence in the inferred ranking. To demonstrate the utility of this technique in evaluating ecological data, we test web-building spiders for predation on a primary prey item, springtails. Using these approaches we found that an orb-weaving spider consumes springtail prey at a higher rate than a syntopic sheet-weaving spider, despite occupying microhabitats where springtails are less frequently encountered. We suggest that spider-web architecture (orb web vs. sheet web) is a primary determinant of prey-consumption rates within this assemblage of predators, which demonstrates the potential influence of predator foraging behaviour on trophic web structure. We also discuss how additional assumptions can be incorporated into the same analysis to allow broader application of the technique beyond the specific example presented. We believe that such modelling techniques can greatly advance the field of molecular gut-content analysis.

Influence of Alternative Prey on the Functional Response of a Predator in Two Contexts: With and without Intraguild Predation.

In biological control, joint releases of predators and parasitoids are standard. However, intraguild predation (IGP) can occur when a predator attacks a parasitoid, potentially affecting pest control dynamics. In addition to the focal prey (FP), Trialeurodes vaporariorum, the intraguild predator (IG-predator) Geocoris punctipes can consume the parasitoid Eretmocerus eremicus (IG-prey). In this IGP context with multiple prey, an alternative prey (AP), like the aphid Myzus persicae, may influence interactions. Theory predicts that, in simple interactions, a predator's functional response (FR) to the FP changes with the presence of an AP. However, whether this holds in an IGP context is unknown. In this study, we empirically tested that prediction. Our results show that without IGP, G. punctipes exhibits a generalized FR with and without AP. Nevertheless, with IGP, the predator exhibited a Type II FR at low and high AP densities, increasing pressure on the FP and potentially favoring short-term biological control strategies. However, when 25 AP were offered, the predator's response shifted, underscoring the importance of monitoring AP densities to prevent potential disruptions in FP control. In both contexts, the increase in AP produced a handling time increase and a decrease in consumption rate. These results indicate that the theoretical prediction of the effect of AP on the FR is met only under specific conditions, and the complexity of multitrophic interactions must be considered.

Trophic omnivory across a productivity gradient: intraguild predation theory and the structure and strength of species interactions.

Intraguild predation theory centres on two predictions: (i) for an omnivore and an intermediate predator (IG-prey) to coexist on shared resources, the IG-prey must be the superior resource competitor, and (ii) increasing resource productivity causes the IG-prey's equilibrium abundance to decline. I tested these predictions with a series of species-rich food webs along New Zealand's rocky shores, focusing on two predatory whelks, Haustrum haustorium, a trophic omnivore, and Haustrum scobina, the IG-prey. In contrast to theory, the IG-prey's abundance increased with productivity. Furthermore, feeding rates and allometric considerations indicate a competitive advantage for the omnivore when non-shared prey are considered, despite the IG-prey's superiority for shared prey. Nevertheless, clear and regular cross-gradient changes in network structure and interaction strengths were observed that challenge the assumptions of current theory. These insights suggest that the consideration of consumer-dependent functional responses, non-equilibrium dynamics, the dynamic nature of prey choice and non-trophic interactions among basal prey will be fruitful avenues for theoretical development.

Indirect food web interactions mediated by predator-rodent dynamics: relative roles of lemmings and voles.

Production cycles in birds are proposed as prime cases of indirect interactions in food webs. They are thought to be driven by predators switching from rodents to bird nests in the crash phase of rodent population cycles. Although rodent cycles are geographically widespread and found in different rodent taxa, bird production cycles appear to be most profound in the high Arctic where lemmings dominate. We hypothesized that this may be due to arctic lemmings inducing stronger predator responses than boreal voles. We tested this hypothesis by estimating predation rates in dummy bird nests during a rodent cycle in low-Arctic tundra. Here, the rodent community consists of a spatially variable mix of one lemming (Lemmus lemmus) and two vole species (Myodes rufocanus and Microtus oeconomus) with similar abundances. In consistence with our hypothesis, lemming peak abundances predicted well crash-phase nest predation rates, whereas the vole abundances had no predictive ability. Corvids were found to be the most important nest predators. Lemmings appear to be accessible to the whole predator community which makes them particularly powerful drivers of food web dynamics.

A comparison of candidate banker plants for management of pests in lettuce.

Agricultural systems are often lacking in resources for natural enemies. Providing alternative prey can help natural enemies persist through periods of low pest abundance, although this approach has been rarely commercially implemented in open field crops. In this study, we tested the potential of eight plant species to provide alternative prey to natural enemies in lettuce fields over a 2-yr period. Results showed that the tested plants would not act as sources of the lettuce aphid Nasonovia ribisnigri Mosley (Hemiptera: Aphididae), the primary lettuce pest. Of the banker plants tested, barley contained high numbers of non-lettuce aphids and appeared to provide reliable habitat for hoverfly larvae. However, lettuce aphids were present on lettuce early in the season, and may have dwarfed any effects of nonlettuce aphids on natural enemy populations. Numbers of hoverfly larvae were also high in lettuce, but did not appear to track numbers of non-lettuce aphids on banker plants. In contrast, numbers of lacewing larvae were highest on plants containing high numbers of non-lettuce aphids, and predatory hemipterans appeared to be associated with numbers of thrips on banker plants. Although barley showed promise as a source of alternative aphids, it did not appear to improve pest control in the adjacent crop.

Pine marten predation of common goldeneye nests: Effects of cavity age and habitat override any effect of microtine rodent abundance.

According to the alternative prey hypothesis (APH), the temporal synchrony in population fluctuations of microtine rodents and other small herbivores in boreal areas is caused by generalist predators with numerical and functional response to microtines, leading to an increased predation of prey alternative to microtines in the low phase of the microtine population fluctuations. The tree-climbing pine marten (Martes martes) is a food generalist that includes bird eggs among its alternative prey, also eggs of the cavity-nesting common goldeneye (Bucephala clangula). We used long-term data to test whether pine marten predation of goldeneye eggs in nest boxes varied as predicted by the APH. As a measure of microtine abundance at the time of nesting, we applied two measures. First, for goldeneye nests located <40 km from our microtine trapping site, we applied the trapping index of microtine rodents. Second, to also use data from nests located >40 km from our microtine trapping site, and from nests in years when trapping was not conducted, we used two proxies for the microtine abundance: whether boreal owls (Aegolius funereus) nested in any of our boxes <40 km from each goldeneye nest and the average clutch size of these boreal owls. The probability of predation of a goldeneye nest was independent of the microtine trapping index and independent of the proxies for microtine abundance. However, it increased with cavity age, taken as the number of nesting seasons elapsed since the actual nest box was installed, and declined with distance from habitat with forest canopy. The effect of cavity age confirms that the long-term spatial memory of pine marten is an important factor in the pattern of its predation on nests in tree cavities.

The abundance of small mammals is positively linked to survival from nest depredation but negatively linked to local recruitment of a ground nesting precocial bird.

Generalist predators using small mammals as their primary prey are suggested to shift hunting alternative prey such as bird nests, when small mammals are in short supply (the alternative prey hypothesis, APH). Nest survival and survival of young individuals should be positively linked to small mammal abundance and negatively linked to predator abundance, but little information exists from survival of chicks, especially until recruitment. We test these predictions of the APH using 13 years (2002-2014) of life history data from a ground nesting shorebird breeding on coastal meadows. We use small mammal abundance in the previous autumn as a proxy for spring predator abundance, mainly of mammalian predators. We examine whether small mammal abundance in the spring and previous autumn explain annual variation in nest survival from depredation and local recruitment of the southern dunlin Calidris alpina schinzii. As predicted by the APH, survival from nest predation was positively linked to spring small mammal abundance and negatively linked to autumn small mammal abundance. Importantly, local recruitment showed opposite responses. This counterintuitive result may be explained by density-dependent survival. When nest depredation rates are low, predators may show stronger numerical and functional responses to high shorebird chick abundance on coastal meadows, whereas in years of high nest depredation, few hatching chicks lure fewer predators. The opposite effects on nest and local recruitment demonstrate the diverse mechanisms by which population size variation in primary prey can affect dynamics of alternative prey populations.

Alternative prey and farming system mediate predation of Colorado potato beetles by generalists.

BACKGROUND: Biological control by generalist predators can be mediated by the abundance and biodiversity of alternative prey. When alternative prey draw predator attacks away from the control target, they can weaken pest suppression. In other cases, a diverse prey base can promote predator abundance and biodiversity, reduce predator-predator interference, and benefit biocontrol. Here, we used molecular gut-content analysis to assess how community composition altered predation of Colorado potato beetle (Leptinotarsa decemlineata (Say)) by Nabis sp. and Geocoris sp. Predators were collected from organic or conventional potato (Solanum tuberosum L.) fields, encouraging differences in arthropod community composition. RESULTS: In organic fields, Nabis predation of potato beetles decreased with increasing arthropod richness and predator abundance. This is consistent with Nabis predators switching to other prey species when available and with growing predator-predator interference. In conventional fields these patterns were reversed, however, with potato beetle predation by Nabis increasing with greater arthropod richness and predator abundance. For Geocoris, Colorado potato beetle predation was more frequent in organic than in conventional fields. However, Geocoris predation of beetles was less frequent in fields with higher abundance of the detritus-feeding fly Scaptomyza pallida Zetterstedt, or of all arthropods, consistent with predators choosing other prey when available. CONCLUSION: Alternative prey generally dampened predation of potato beetles, suggesting these pests were less-preferred prey. Nabis and Geocoris differed in which alternative prey were most disruptive to feeding on potato beetles, and in the effects of farm management on predation, consistent with the two predator species occupying complementary feeding niches. © 2021 Society of Chemical Industry.

Prey choice by a freshwater copepod on larval Aedes mosquitoes in the presence of alternative prey.

Predator-prey interactions can have a significant impact on the abundance and distribution of species, but the outcome of these interactions is often context-dependent. In small freshwater habitats, predacious copepods are potential biological control agents for mosquito larvae. Through laboratory experiments, we tested if the presence of a non-mosquito prey (neonate Daphnia pulex) influenced prey selection of the predaceous copepod (Acanthocyclops vernalis) on 1st instar Aedes mosquitoes (Aedes aegypti and Aedes albopictus). Copepods were starved for 12 h prior to being exposed to the two prey types (larval mosquitoes and Daphnia) at three densities: 25 mosquitoes:75 Daphnia, 50 mosquitoes:50 Daphnia, 75 mosquitoes:25 Daphnia. Single prey choice trials for each species as well as no-predator trials were also established for controls. Copepods were effective predators, with a single copepod consuming up to 37 1st instar mosquito larvae during the 24-h trials. The number of mosquitoes consumed increased with their relative density, but the proportion of mosquitoes consumed was highest when Aedes made up only 25% of the population. Results from our study show that in a simple predator/two-prey system, two species of larval mosquitoes (Ae. aegypti and Ae. albopictus) are preferentially consumed over an alternative zooplankton by the copepod predator Acanthocyclops vernalis.

Establishment of a Faba Bean Banker Plant System with Predator Orius strigicollis for the Control of Thrips Dendrothrips minowai on Tea Plants under Laboratory Conditions.

The stick tea thrip Dendrothrips minowai (Priesner) (Thysanoptera: Thripidae) is a destructive pest in tea plantations in south and southwest China. To control this pest, a non-crop banker plant system was developed using a polyphagous predator Orius strigicollis (Poppius) (Heteroptera: Anthocoridae) with the black bean aphid Aphis fabae (Scopoli) (Hemiptera: Aphididae) as an alternative prey and the faba bean Vicia faba as the banker plant to support the predator in targeting the pest. The fitness of A. fabae on tea plants and faba bean was evaluated to determine its host specificity. Moreover, the control efficacy of the banker plant system on D. minowai on tea plants was tested in the laboratory and compared with that of direct release of O. strigicollis. The experiments showed that faba bean was an excellent non-crop host for A. fabae because, while the aphid population increased quickly on faba bean, it could only survive for up to 9 days on tea plants. Compared with direct release of O. strigicollis, lower densities of pest were observed when introducing the banker plant system. Our results indicate that this banker plant system has the potential to be implemented in the field to improve the control of the pest thrips.

Dinophysis Ehrenberg (Dinophyceae) in Southern Chile harbours red cryptophyte plastids from Rhodomonas/Storeatula clade.

Photosynthetic species of the dinoflagellate genus Dinophysis are known to retain temporary cryptophyte plastids of the Teleaulax/Plagioselmis/Geminigera clade after feeding the ciliate Mesodinium rubrum. In the present study, partial plastid 23S rDNA sequences were retrieved in Southern Chilean waters from oceanic (Los Lagos region), and fjord systems (Aysén region), in single cells of Dinophysis and accompanying organisms (the heliozoan Actinophrys cf. sol and tintinnid ciliates), identified by means of morphological discrimination under the light microscope. All plastid 23S rDNA sequences (n = 23) from Dinophysis spp. (Dinophysis acuta, D. caudata, D. tripos and D. subcircularis) belonged to cryptophytes from clade V (Rhinomonas, Rhodomonas and Storeatula), although they could not be identified at genus level. Moreover, five plastid sequences obtained from heliozoans (Actinophryida, tentatively identified as Actinophrys cf. sol), and tintinnid ciliates, grouped together with those cryptophyte sequences. In contrast, two additional sequences from tintinnids belonged to other taxa (chlorophytes and cyanobacteria). Overall, the present study represents the first time that red cryptophyte plastids outside of the Teleaulax/Plagioselmis/Geminigera clade dominate in wild photosynthetic Dinophysis spp. These findings suggest that either Dinophysis spp. are able to feed on other ciliate prey than Mesodinium and/or that cryptophyte plastids from clade V prevail in members of the M. rubrum species complex in the studied area.

Implications of shared predation for space use in two sympatric leporids.

Spatial variation in habitat riskiness has a major influence on the predator-prey space race. However, the outcome of this race can be modulated if prey shares enemies with fellow prey (i.e., another prey species). Sharing of natural enemies may result in apparent competition, and its implications for prey space use remain poorly studied. Our objective was to test how prey species spend time among habitats that differ in riskiness, and how shared predation modulates the space use by prey species. We studied a one-predator, two-prey system in a coastal dune landscape in the Netherlands with the European hare (Lepus europaeus) and European rabbit (Oryctolagus cuniculus) as sympatric prey species and red fox (Vulpes vulpes) as their main predator. The fine-scale space use by each species was quantified using camera traps. We quantified residence time as an index of space use. Hares and rabbits spent time differently among habitats that differ in riskiness. Space use by predators and habitat riskiness affected space use by hares more strongly than space use by rabbits. Residence time of hare was shorter in habitats in which the predator was efficient in searching or capturing prey species. However, hares spent more time in edge habitat when foxes were present, even though foxes are considered ambush predators. Shared predation affected the predator-prey space race for hares positively, and more strongly than the predator-prey space race for rabbits, which were not affected. Shared predation reversed the predator-prey space race between foxes and hares, whereas shared predation possibly also released a negative association and promoted a positive association between our two sympatric prey species. Habitat riskiness, species presence, and prey species' escape mode and foraging mode (i.e., central-place vs. noncentral-place forager) affected the prey space race under shared predation.

Crustacean amphipods from marsh ponds: a nutritious feed resource with potential for application in Integrated Multi-Trophic Aquaculture.

Coastal protection, nutrient cycling, erosion control, water purification, and carbon sequestration are ecosystem services provided by salt marshes. Additionally, salt ponds offer coastal breeding and a nursery habitat for fishes and they provide abundant invertebrates, such as amphipods, which are potentially useful as a resource in aquaculture. Fishmeal and fish oil are necessary food resources to support aquaculture of carnivorous species due to their omega-3 long-chain polyunsaturated fatty acids (n-3 LC-PUFA). Currently, aquaculture depends on limited fisheries and feed with elevated n-3 LC-PUFA levels, but the development of more sustainable food sources is necessary. Amphipods appear to be a potential high quality alternative feed resource for aquaculture. Hence, a nutritional study was carried out for several main amphipod species-Microdeutopus gryllotalpa, Monocorophium acherusicum, Gammarus insensibilis, Melita palmata and Cymadusa filosa-in terrestrial ponds in the South of Spain. These species showed high protein content (up to 40%), high n-3 PUFA and phospholipid levels, and high levels of phophatidylcholine (PC), phosphatidylethanolamine (PE) and triacylglycerols (TAG), the latter being significantly high for M. acherusicum. M. gryllotalpa and M. acherusicum showed the highest proportion of lipids (19.15% and 18.35%, respectively). Isoleucine, glycine and alanine were the dominant amino acids in all species. In addition, amphipods collected from ponds showed low levels of heavy metals. Furthermore, the biochemical profiles of the five species of amphipods have been compared with other studied alternative prey. Therefore, pond amphipods are good candidates to be used as feed, and are proposed as a new sustainable economic resource to be used in aquaculture. G. insensibilis may be the best for intensive culture as an alternative feed resource because it shows: (1) adequate n-3 PUFA and PL composition; (2) high levels of glycine, alanine, tyrosine, isoleucine and lysine; (3) high natural densities; (4) large body size (≥1 cm), and (5) high concentration of calcium. Moreover, a combined culture of amphipods and fishes in these marsh ponds seems a promising and environmentally sustainable way to develop Integrate Multi-Trophic Aquaculture (IMTA) in these ecosystems.