Does prey preference change as a result of prey species being presented together? Analysis of prey selection by the predatory mite Typhlodromus pyri (Acarina: Phytoseiidae)

Summary Prey-selection behaviour of the phytoseiid mite Typhlodromus pyri Scheuten was analysed with a Markovtype model of feeding-state dynamics and feeding-state dependent searching behaviour (Sabelis 1981, 1986, 1989; Metz and Van Batenburg 1985a, b). All behavioural characteristics of the predator which are independent of the feeding state were represented by one parameter. The remaining feeding-state dependent characteristics were represented by a function of the feeding state, with one parameter. The best parameter values to describe a predator-prey interaction were determined by fitting the model to the predation rates in monocultures. Under the assumption that the parameter values are not dependent on the composition of prey species supply, the diet of the predators in mixed cultures was predicted from parameters estimated in monoculture experiments.Two prey types, apple rust mite (Aculus schlechtendali (Nalepa)) adults and European red spider mite (Panonychus ulmi (Koch)) larvae were studied. A large discrepancy was observed between calculated and experimentally determined predation rates of T. pyri in mixed cultures: the predators actually killed 3–7 times more P. ulmi larvae than was predicted by the model.The large difference between observed and predicted predation rates in mixed cultures cannot be explained by changes in the behaviour of the prey species as a result of being together. Therefore, it seems likely that the prey selection behaviour of the predator was different when prey species were presented together than when presented singly. Apparently the predatory mite T. pyri prefers P. ulmi to S. schlechtendali.     

Does prescribed burning result in biotic homogenization of coastal heathlands?

Biotic homogenization due to replacement of native biodiversity by widespread generalist species has been demonstrated in a number of ecosystems and taxonomic groups worldwide, causing growing conservation concern. Human disturbance is a key driver of biotic homogenization, suggesting potential conservation challenges in seminatural ecosystems, where anthropogenic disturbances such as grazing and burning are necessary for maintaining ecological dynamics and functioning. We test whether prescribed burning results in biotic homogenization in the coastal heathlands of north‐western Europe, a seminatural landscape where extensive grazing and burning has constituted the traditional land‐use practice over the past 6000 years. We compare the beta‐diversity before and after fire at three ecological scales: within local vegetation patches, between wet and dry heathland patches within landscapes, and along a 470 km bioclimatic gradient. Within local patches, we found no evidence of homogenization after fire; species richness increased, and the species that entered the burnt Calluna stands were not widespread specialists but native grasses and herbs characteristic of the heathland system. At the landscapes scale, we saw a weak homogenization as wet and dry heathland patches become more compositionally similar after fire. This was because of a decrease in habitat‐specific species unique to either wet or dry habitats and postfire colonization by a set of heathland specialists that established in both habitat types. Along the bioclimatic gradient, species that increased after fire generally had more specific environmental requirements and narrower geographical distributions than the prefire flora, resulting in a biotic ‘heterogenisation’ after fire. Our study demonstrates that human disturbance does not necessarily cause biotic homogenization, but that continuation of traditional land‐use practices can instead be crucial for the maintenance of the diversity and ecological function of a seminatural ecosystem. The species that established after prescribed burning were heathland specialists with relatively narrow geographical ranges.     

Does follicle excision always result in enlargement of offspring size in lizards?

An experimental reduction of offspring number has been reported to result in enlargement of offspring size in lizards. We applied the “follicle excision” technique to a lacertid lizard (Takydromus septentrionalis) to examine whether this effect is generalisable to lizards. Of the 82 females that produced 3 successive clutches in the laboratory, 23 females underwent follicle excision after they oviposited the first clutch. Follicle excision reduced clutch size, but did not alter egg size. This result indicates that egg size is not altered during vitellogenesis in T. septentrionalis. Females undergoing follicle excision produced a third clutch (a second post-surgical clutch) as normally as did control females. Females switched from producing more but smaller eggs early in the breeding season to fewer but larger eggs later in the season. Our results indicate that female T. septentrionalis maximize reproductive success by diverting an optimal, rather than a higher, fraction of the available energy to individual offspring. This optimized allocation of the available energy to offspring production explains why follicle excision does not result in enlargement of egg size in this species. Our study provides evidence that an experimental reduction of offspring number does not always result in enlargement of offspring size in lizards.     

Does dust result in mite outbreaks in apple orchards?

The effect of dust on phytophagous mite numbers was examined in five apple orchards situated in the dry, inland apple producing Ceres area, South Africa. The study was conducted over three seasons. The season with the most dust had the least number of mites. There was no relationship between the amount of dust on leaves and mite numbers from different orchards. Of the 15 correlations between the amount of dust on individual trees and the number of mites on these trees, two were marginally not significant and one was highly significant, but negative. Therefore, seasons during which there is a lot of dust did not result in mite outbreaks nor did dusty orchards harbour elevated mite population levels, and trees with a lot of dust did not necessarily harbour more mites than trees with less dust. However, if there is enough dust to cause stress to the trees, phytophagous mite outbreaks could occur.     

Does elevated testosterone result in increased exposure and transmission of parasites?

Male-biased infection is a common phenomenon in vertebrate-parasite systems and male-biased transmission has been experimentally demonstrated. One mechanism that is hypothesized to create male-biased transmission is the immuno-suppressive effect of testosterone because it increases susceptibility to infection. Testosterone also influences host behaviour and, consequently, may increase exposure to parasites. To test how testosterone could increase exposure and transmission, we undertook a longitudinal mark-recapture study where we experimentally elevated testosterone levels in wild male rodents. Individuals in control populations reduced the average number of contacts over the treatment period, while populations with experimentally elevated testosterone levels maintained the number of contacts between hosts. As a result, the transmission potential was higher in testosterone treated populations compared to controls. Our results indicated that males with high-testosterone levels alter the population-level contacts, producing different social networks and increasing transmission potential compared to those where testosterone is at background levels.     

Individual behavioral variation in predator–prey models

The role of individual behavioral variation in community dynamics was studied. Behavioral variation in this study does not refer to differences in average responses (e.g., average response between presence and absence of antipredator behavior). Rather it refers to the variation around the average response that is not explained by trivial experimental treatments. First, the effect of behavioral variation was examined based on Jensen’s inequality. In cases of commonly used modeling framework with type II functional response, neglecting behavioral variation (a component of encounter rate) causes overestimation of predation effects. The effect of this bias on community processes was examined by incorporating the behavioral variation in a commonly used consumer-resource model (Rosenzweig–MacArthur model). How such a consideration affects a model prediction (paradox of enrichment) was examined. The inclusion of behavioral variation can both quantitatively and qualitatively alter the model characteristics. Behavioral variation can substantially increase the stability of the community with respect to enrichment.     

Does sun glare increase antipredator behaviour in prey?

As the sun gradually lowers over the horizon, prey species with more sun in their eyes should have more difficulty in visually monitoring their surroundings for threats and thus experience a higher predation risk. In a unique setting, I could examine changes in antipredator behaviour in a prey species, the semipalmated sandpiper Calidris pusilla, facing attacks by peregrine falcons Falco peregrinus, which originated from the general direction of the lowering sun. I predicted gradual changes in antipredator behaviour as sun glare becomes more problematic later in the day. As the day progressed, sandpipers occurred in sparser groups when the sun glared but not when clouds obscured the sun, suggesting that fewer individuals engaged in risky foraging. Pecking rate and foraging success decreased later in the day when the sun glared but not otherwise implying an increase in vigilance at the expense of foraging. When more sun hit their eyes, sandpipers also moved faster suggesting increased skittishness. The sun glare effect might be relevant to any species foraging in open areas not only when the sun sets but also when it rises especially if predators can target prey species at these vulnerable times. The temporal gradient in predation risk that the sun glare effect creates might thus apply broadly and have important consequences for antipredator vigilance, foraging efficiency, and habitat use.     

Does colour polymorphism enhance survival of prey populations?

That colour polymorphism may protect prey populations from predation is an old but rarely tested hypothesis. We examine whether colour polymorphic populations of prey exposed to avian predators in an ecologically valid visual context were exposed to increased extinction risk compared with monomorphic populations. We made 2976 artificial pastry prey, resembling Lepidoptera larvae, in four different colours and presented them in 124 monomorphic and 124 tetramorphic populations on tree trunks and branches such that they would be exposed to predation by free-living birds, and monitored their ‘survival’. Among monomorphic populations, there was a significant effect of prey coloration on survival, confirming that coloration influenced susceptibility to visually oriented predators. Survival of polymorphic populations was inferior to that of monomorphic green populations, but did not differ significantly from monomorphic brown, yellow or red populations. Differences in survival within polymorphic populations paralleled those seen among monomorphic populations; the red morph most frequently went extinct first and the green morph most frequently survived the longest. Our findings do not support the traditional protective polymorphism hypothesis and are in conflict with those of earlier studies. As a possible explanation to our findings, we offer a competing ‘giveaway cue’ hypothesis: that polymorphic populations may include one morph that attracts the attention of predators and that polymorphic populations therefore may suffer increased predation compared with some monomorphic populations.     

Signal convergence in fruits: a result of selection by frugivores?

The Dispersal Syndrome hypothesis remains contentious, stating that apparently nonrandom associations of fruit characteristics result from selection by seed dispersers. We examine a key assumption under this hypothesis, i.e. that fruit traits can be used as reliable signals by frugivores. We first test this assumption by looking at whether fruit colour allows birds and primates to distinguish between fruits commonly dispersed by birds or primates. Second, we test whether the colours of fruits dispersed by primates are more contrasting to primates than the colours of bird‐dispersed fruits, expected if fruit colour is an adaptation to facilitate the detection by seed dispersers. Third, we test whether fruit colour has converged in unrelated plant species dispersed by similar frugivores. We use vision models based on peak sensitivities of birds’ and primates’ cone cells. We base our analyses on the visual systems of two types of birds (violet and ultraviolet based) and three types of primates (trichromatic primates from the Old and the New Worlds, and a dichromatic New World monkey). Using a Discriminant Function Analysis, we find that all frugivore groups can reliably discriminate between bird‐ and primate‐dispersed fruits. Fruit colour can be a reliable signal to different seed dispersers. However, the colours of primate‐dispersed fruits are less contrasting to primates than those of bird‐dispersed fruits. Fruit colour convergence in unrelated plants is independent of phylogeny and can be better explained by disperser type, which supports the hypothesis that frugivores are important in fruit evolution. We discuss adaptive and nonadaptive hypotheses that can potentially explain the pattern we found.     

Does intraspecific niche partitioning in a native predator influence its response to an invasion by a toxic prey species?

Abstract The introduced and highly toxic cane toad (Bufo marinus) is rapidly spreading across northern Australia where it may affect populations of large terrestrial vertebrate predators. The ecological impact of cane toads will depend upon the diets, foraging modes and habitat use of native predators, and their feeding responses to cane toads. However, intraspecific niche partitioning may influence the degree of vulnerability of predators to toxic prey, as well as the time course of the impact of alien invaders on native species. We studied the diet of the northern death adder Acanthophis praelongus and their feeding responses to cane toads. In the laboratory, death adders from all size classes and sexes readily consumed frogs and cane toads. Diets of free ranging A. praelongus from the Adelaide River floodplain were more heterogeneous. Juvenile snakes ate mainly frogs (39% of prey items) and small scincid lizards (43%). Both sexes displayed an ontogenetic dietary shift from lizards to mammals, but adult males fed on frogs (49%) and mammals (39%) whereas adult females (which grew larger than males) fed mainly on mammals (91%) and occasionally, frogs (9%). Feeding rates and body condition of adult snakes varied temporally and tracked fluctuations in prey availability. These results suggest that cane toads may negatively affect populations of northern death adders in the Darwin region. However, we predict that different size and sex classes of A. praelongus will experience differential mortality rates over different timescales. The initial invasion of large toads may affect adult males, but juveniles may be unaffected until juvenile toads appear the following year, and major affects on adult female death adders may be delayed until annual rainfall fluctuations reduce the availability of alternative (rodent) prey.     

Does diet in Celtic Sea fishes reflect prey availability?

Feeding preferences of Celtic Sea fishes were investigated using a database of stomach content records, collected between 1977 and 1994. The diet of cod Gadus morhua , hake Merluccius merluccius , megrim Lepidorhombus whiffiagonis , whiting Merlangius merlangus and saithe Pollachius virens changed markedly as the animals grew larger, and although large predators generally chose larger bodied prey, the variability of prey sizes consumed also increased. Large predators continued to select small, low value, benthic prey ( e.g . Callionymus spp. and Trisopterus spp.) which were easier to catch, rather than larger, more energy lucrative pelagic prey ( e.g . mackerel Scomber scombrus ), even though these pelagic prey‐fishes were nearly always available and were often very abundant. Stock estimates of the International Council for the Exploration of the Sea and U.K. groundfish survey catches were used as indices of prey abundance. Blue‐whiting Micromesistius poutassou and other small pelagic fishes ( Argentina spp. and clupeoids) were identified as being particularly important, and were consumed by some predators more often than would be expected given the abundance of these prey in the environment. There was no evidence for density‐dependent feeding by predators on mackerel and only hake exhibited density‐dependent feeding on horse‐mackerel. Hake, cod and megrim consumed more blue‐whiting when this prey was at higher abundance in the environment. In choosing what prey to consume, predators must balance costs and benefits, considering the quality of prey and the energy expended during search, capture and handling.     

Does prey community composition affect the way different behavioral types interact with their environment?

We examined how different exploratory behavioral types of largemouth bass responded to differing prey communities by determining effects on growth, survival and diet in experimental ponds. We found evidence that non-explorer largemouth bass target young-of-year bluegill early on in life, but bluegill were not an important diet item by late summer. The presence of young-of-year bluegill as prey does appear to affect the foraging strategy of the two exploring types differently. In the absence of small bluegill, both behavioral types feed primarily on benthic invertebrates and zooplankton. When small bluegill were present, we saw a shift away from zooplankton as prey for largemouth bass. However, that shift was toward more benthic invertebrates for non-exploring behavioral types and toward terrestrial insects for exploring behavioral types. Thus, it appears that prey community composition can have important effects on the way in which different behavioral types interact with their environment.     

Are behavioural traits in prey sensitive to the risk imposed by predatory fish?

1. Behavioural differences among prey species may result from evolutionary adaptations that facilitate coexistence with different predators and influence vulnerability to predators. It has been hypothesised that prey species modify their behaviour in relation to the risk posed by particular predators. 2. We examined the relationship between anti‐predator behaviour and predation risk in five species of larval odonates in combination with three predatory fish species (perch, gudgeon and rudd) that differ in foraging behaviour. The odonates, Platycnemis pennipes, Coenagrion puella, Lestes sponsa, Sympetrum striolatum and Libellula depressa, differ with regard to their life cycle and habitat, including water depth, occurrence in temporary ponds and co‐existence with fish. 3. The odonate species differed in their response to fish: (i) Two species showed a flexible response. Larval C. puella reduced activity in the presence of fish, regardless of species, whereas L. depressa altered their activity only in the presence of gudgeon. (ii) Independent of fish species, all odonates except L. depressa exhibited spatial avoidance of fish. This was interpreted as a more general anti‐predator response. (iii) In some cases the odonates showed no response to predators and their behaviour was thus independent of predation risk. 4. Our results confirm that all odonates responded to the presence of at least some predatory fish, and that some odonate species discriminated between fish species. However, we found no significant correlation between behavioural modifications and predation risk, indicating that anti‐predator responses and predation risk depend on the particular predator and the species being preyed on.     

Does local adaptation to resources explain genetic differentiation among Daphnia populations?

Substantial genetic differentiation is frequently observed among populations of cyclically parthenogenetic zooplankton despite their high dispersal capabilities and potential for gene flow. Local adaptation has been invoked to explain population genetic differentiation despite high dispersal, but several neutral models that account for basic life history features also predict high genetic differentiation. Here, we study genetic differentiation among four populations of Daphnia pulex in east central Illinois. As with other studies of Daphnia, we demonstrate substantial population genetic differentiation despite close geographic proximity (<50 km; mean θ = 0.22). However, we explicitly tested and failed to find evidence for, the hypothesis that local adaptation to food resources occurs in these populations. Recognizing that local adaptation can occur in traits unrelated to resources, we estimated contemporary migration rates (m) and tested for admixture to evaluate the hypothesis that observed genetic differentiation is consistent with local adaptation to other untested ecological factors. Using Bayesian assignment methods, we detected migrants in three of the four study populations including substantial evidence for successful reproduction by immigrants in one pond, allowing us to reject the hypothesis that local adaptation limits gene flow for at least this population. Thus, we suggest that local adaptation does not explain genetic differentiation among these Daphnia populations and that other factors related to extinction/colonization dynamics, a long approach to equilibrium F_ST or substantial genetic drift due to a low number of individuals hatching from the egg bank each season may explain genetic differentiation.     

Selection of aphid prey by a generalist predator: do prey chemical defences matter?

1. For predators, prey selection should maximise nutrition and minimise fitness costs. In the present study, it was investigated whether a generalist predator [Chrysoperla carnea (Stephens) lacewing larvae] rejected harmful, chemically‐defended prey [Brevicoryne brassicae (Linnaeus) aphids] when non‐defended prey [Myzus persicae (Sulzer) aphids] were available. 2. It was tested: (i) whether consuming different prey species affects predator mortality; (ii) whether naïve predators reject chemically‐defended prey while foraging when non‐defended prey are available; (iii) whether the relative abundance of each prey affects the predator's prey choice; and (iv) whether predators learn to avoid consuming chemically‐defended prey after exposure to both prey species. 3. Consumption of B. brassicae yielded greater C. carnea mortality than M. persicae consumption, but naïve C. carnea did not reject B. brassicae in favour of M. persicae during foraging. When presented at unequal abundances, naïve predators generally consumed each aphid species according to their initial relative abundance, although, predation of non‐defended prey was less than expected when defended prey were initially more abundant, indicating a high consumption of B. brassicae impeded M. persicae consumption. With experience, C. carnea maintained predation of both aphid species but consumed more M. persicae than B. brassicae, indicating a change in behaviour. 4. Although prey choice by C. carnea may change with experience of available prey, prey chemical defences do not appear to influence prey choice by naïve predators. This inability to avoid harmful prey could facilitate wider, indirect interactions. Myzus persicae may benefit where high consumption of B. brassicae hinders predators in the short term, and in the long term, increases predator mortality.     

Phenotypic shifts caused by predation: selection or life-history shifts?

Predators can impose both selection and life-history shifts in prey populations. Because both processes may affect phenotypic distributions, the estimates of selection differentials may be biased. We carried out two field experiments to disentangle these separate effects. We studied whether dragonfly predation by Aeshna cyanea changes the distributions in body size and lamellae morphology in the damselfly Lestes sponsa. Damselflies have caudal lamellae which are used in escapes by swimming. In a first experiment, we manipulated predator presence (No Aeshna, Encaged Aeshna or Free-ranging Aeshna) and stopped the experiment when all larvae had moulted once. In a second experiment, larvae were confronted with a Free-ranging Aeshna but collected before moulting, and survivors were compared with a control sample taken at the start of the experiment. The presence of Aeshna largely reduced the survival probabilities of the Lestes larvae at a very similar rate in both experiments. Daily survival probabilities did not differ between the No Aeshna and Encaged Aeshna treatments. In the Free-ranging Aeshna treatment of the first experiment, size was reduced compared to the other two treatments, creating a significant apparent selection differential. This was probably mainly due to predator-induced reduced growth because in the second experiment, where growth effects were excluded, size of the survivors did not differ from the control sample. In both experiments there was a significant selection pressure for larger lamellae. Standardized directional selection differentials were similar in both experiments (0.57 and 0.28 phenotypic standard deviation units). No survival selection on lamellae shape was detected. These results are in agreement with previous findings that lamellae size, but not lamellae shape, enhances swimming performance and thereby predator escape in this species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Does migration of hybrids contribute to post-zygotic isolation in flycatchers?

In the face of hybridization, species integrity can only be maintained through post-zygotic isolating barriers (PIBs). PIBs need not only be intrinsic (i.e. hybrid inviability and sterility caused by developmental incompatibilities), but also can be extrinsic due to the hybrid's intermediate phenotype falling between the parental niches. For example, in migratory species, hybrid fitness might be reduced as a result of intermediate migration pathways and reaching suboptimal wintering grounds. Here, we test this idea by comparing the juvenile to adult survival probabilities as well as the wintering grounds of pied flycatchers (Ficedula hypoleuca), collared flycatchers (Ficedula albicollis) and their hybrids using stable isotope ratios of carbon (delta13C) and nitrogen (delta15N) in feathers developed at the wintering site. Our result supports earlier observations of largely segregated wintering grounds of the two parental species. The isotope signature of hybrids clustered with that of pied flycatchers. We argue that this pattern can explain the high annual survival of hybrid flycatchers. Hence, dominant expression of the traits of one of the parental species in hybrids may substantially reduce the ecological costs of hybridization.     

Are behavioral responses to eyespots in sticklebacks influenced by the visual environment? An experimental examination

Eyespots are taxonomically widespread color patterns consisting of large concentric rings that are commonly assumed to protect prey by influencing the behaviors of predators. Although there is ample experimental evidence supporting an anti‐predator function of eyespots in terrestrial animals, whether eyespots have a similar deterring function in aquatic animals remains unclear. Furthermore, studies in terrestrial systems suggest that the protective function of eyespots depends on ambient light conditions where predators encounter them, but this effect has never been tested in aquatic environments. Here, we examine how eyespots influence behavioral responses in an aquatic environment under different visual environments, using laboratory‐reared three‐spined sticklebacks (Gasterosteus aculeatus) as model predators. Specifically, we experimentally examined behavioral responses of sticklebacks toward artificial prey patterns (control vs. eyespots) under two different light environment treatments (low vs. high). We found that eyespots did not postpone attacks from sticklebacks. However, sticklebacks approaching eyespots stopped more frequently than sticklebacks approaching prey items with a control pattern. Sticklebacks were (marginally) slower to attack prey in the low‐light treatment, but the light level did not influence stickleback behavioral responses toward eyespots. We conclude that eyespots can modulate some behaviors of an aquatic predator, albeit with a different functional role from that previously demonstrated in terrestrial species.     

Can photosynthesis respond to short-term fluctuations in atmospheric carbon dioxide?

Rapid and irregular variations of atmospheric CO2 concentrations (ca) occur in nature but are often very much more pronounced and frequent when artificially enriching CO2 concentrations in simulating the future atmosphere. Therefore, there is the danger that plant responses at elevated CO2 in fumigation experiments might reflect the increased frequency and amplitude of fluctuation in concentration as well as the increase in average concentration. Tests were conducted to determine whether the photosynthetic process could sense such fluctuations in ca. Instantaneous chlorophyll fluorescence (Ft) was monitored for wheat leaves (Triticum aestivum cv. Hereward) exposed to ca oscillating symmetrically by 225 mol mol-1 about a ca set point concentration of 575 or 650 mol mol-1. No Ft response was detected to half-cycle step changes in ca lasting less than two seconds, but at half-cycles of two seconds or longer, the response of Ft was pronounced. In order to determine the in vivo linear electron transport rate (J) the O2 concentration was maintained at 21 mmol mol-1 to eliminate photorespiration. J which is directly proportional to the rate of CO2 uptake under these conditions, was not significantly changed at half-cycles of 30 s or less but was decreased by half-cycles of 60 s or longer. It was inferred that if duration of an oscillation is less than 1 minute and is symmetrical with respect to mean CO2 concentration, then there is no effect on current carbon uptake, but oscillations of 1 minute or more decrease photosynthetic CO2 uptake in wheat.