As good as dead? Sublethal predation facilitates lethal predation on an intertidal clam

Although ecologists have speculated that sublethal predation can impact prey dynamics, consequences of these predator effects have seldom been experimentally tested. In soft‐sediment marine communities, fishes crop extended feeding siphons of buried clams, potentially causing clams to reduce their burial depth, thereby enhancing their susceptibility to excavating lethal predators. We simulated cropping of the confamilial clams, Protothaca staminea and Venerupis philippinarum, by removing the top 40% of siphons, which caused each species to burrow 33–50% shallower than conspecifics with intact siphons. To examine subsequent consequences of reduced burial depth, we exposed cropped and intact clams to natural levels of predation in the field. Because of a naturally longer siphon, Protothaca, even after cropping, remained at relatively safe burial depths. In contrast, siphon cropping nearly doubled the mortality rate of Venerupis. Thus, while sublethal predation facilitates lethal predation, this linkage depends on specific life history characteristics, even among ecologically similar species.     

How well do predation rates on artificial nests estimate predation on natural passerine nests?

Daily survival rates (DSR) of three nest types were compared among three passerine species in woodland habitats of the Czech Republic in 1998‐99. The species were: Yellowhammer (YH) Emberiza citrinella, Blackcap (BC) Sylvia atricapilla, Song Thrush (ST) Turdus philomelos. The nest types were: AA, artificial nests with artificial plastic eggs, both mimicking the real models (total n= 432); NA, natural nests left in their original position and baited with artificial plastic eggs (n= 706); and NN, active natural nests observed during the egg stage (n= 596). Rodents were dominant predators of the near‐ground YH nests while predation by corvids slightly prevailed in the shrub nests of BC and ST. There was no consistent relationship between nest type (AA vs. NA) and type of predator. Effects of nest type (AA vs. NA or AA vs. NN, respectively), species, date and study plot on DSR were examined by fitting logistic regression models. Marginally significant (< P < 0.05) differences were found between AA and NA nests in both years. Due to the interaction with date in 1998, the model predicted lower/higher DSR for AA nests before/after about 25 May respectively. In 1999 the DSR of AA nests was consistently higher (difference: 0.011; 95% CI: 0.001 to 0.021). Marginally significant differences between AA and NN nests were found only in 1998: the DSR of AA nests was consistently lower (‐0.014, 95% CI: ‐0.027 to ‐0.001). Comparisons of mean DSR within each species‐year sample showed that the differences between nest types were of both directions and varied from almost zero to 0.02 (ST, BC; ns) or 0.04 (YH; P < 0.05). Although the experimental nests accurately reflected the mean values as well as seasonal trends and interspecific differences in DSR of the real nests in some subsets of data, the pattern was not consistent across species and between years. This study suggests that artificial nests could not be used to assess survival of natural nests without careful validation of the experimental results.     

Vulnerability of black grouse hens to goshawk predation: result of food supply or predation facilitation?

The plant cycle hypothesis says that poor-quality food affects both herbivorous voles (Microtinae spp.) and grouse (Tetraonidae spp.) in vole decline years, leading to increased foraging effort in female grouse and thus a higher risk of predation by the goshawk Accipiter gentilis. Poor-quality food (mainly the bilberry Vaccinium myrtillus) for these herbivores is induced by seed masting failure in the previous year, when the bilberry is able to allocate resources for chemical defence (the mast depression hypothesis; MDH). The predation facilitation hypothesis (PFH) in turn states that increased searching activity of vole-eating predators during or after the decline year of voles disturbs incubating and brooding grouse females. The behaviours used by grouse to avoid these terrestrial predators make them more vulnerable to predation by goshawks. We tested the main predictions of the MDH and PFH by collecting long-term (21-year) data from black grouse Tetrao tetrix hens and cocks killed by breeding goshawks supplemented with indices of bilberry crop, vole abundance and small carnivores in the vicinity of Oulu, northern Finland. We did not find obvious support for the prediction of the MDH that there is a negative correlation of bilberry crop in year t with vole abundance and with predation index of black grouse hens in year t + 1. We did find obvious support for the prediction of the PFH that there is a positive correlation between predator abundance and predation index of grouse hens, because the stoat Mustela erminea abundance index was positively related to the predation index of black grouse hens. We suggest that changes in vulnerability of grouse hens may mainly be caused by the guild of vole-eating predators, who shift to alternative prey in the decline phase of the vole cycle, and thus chase grouse hens and chicks to the talons of goshawks and other avian predators.     

Does predation maintain tit community diversity?

European tits of the genus Parus constitute a complex group of coexisting boreal birds. Here we present a survey of the distribution of three coniferous-living Parus species and one of their main predators, the pygmy owl ( Glaucidium passerinum ), on nine isolated islands in Scandinavia. On all islands the coal tit ( Parus ater ) is the sole tit species when the pygmy owl is absent. The two larger species, the willow tit ( P. montanus ) and the crested tit ( P. cristatus ), only coexist with the coal tit when pygmy owls are present. We suggest that the coexistence of willow tits, crested tits and coal tits is the result of a combination of competition for food and predator-safe foraging sites. The smaller coal tit is superior in exploitation competition for food, while the two larger species have an advantage in interference competition for predator-safe foraging sites. The association between the distribution of the pygmy owl and the two larger tit species on isolated islands in Scandinavia is consistent with the idea that the pygmy owl is a keystone predator.     

Does predation contribute to tree diversity?

Seed and seedling predation may differentially affect competitively superior tree species to increase the relative recruitment success of poor competitors and contribute to the coexistence of tree species. We examined the effect of seed and seedling predation on the seedling recruitment of three tree species, Acer rubrum (red maple), Liriodendron tulipifera (yellow poplar), and Quercus rubra (northern red oak), over three years by manipulating seed and seedling exposure to predators under contrasting microsite conditions of shrub cover, leaf litter, and overstory canopy. Species rankings of seedling emergence were constant across microsites, regardless of exposure to seed predators, but varied across years. A. rubrum had the highest emergence probabilities across microsites in 1997, but Q. rubra had the highest emergence probabilities in 1999. Predators decreased seedling survival uniformly across species, but did not affect relative growth rates (RGRs). Q. rubra had the highest seedling survivorship across microsites, while L. tulipifera had the highest RGRs. Our results suggest that annual variability in recruitment success contributes more to seedling diversity than differential predation across microsites. We synthesized our results from separate seedling emergence and survival experiments to project seedling bank composition. With equal fecundity assumed across species, Q. rubra dominated the seedling bank, capturing 90% of the regeneration sites on average, followed by A. rubrum (8% of sites) and L. tulipifera (2% of sites). When seed abundance was weighted by species-specific fecundity, seedling bank composition was more diverse; L. tulipifera captured 62% of the regeneration sites, followed by A. rubrum (21% of sites) and Q. rubra (17% of sites). Tradeoffs between seedling performance and fecundity may promote the diversity of seedling regeneration by increasing the probability of inferior competitors capturing regeneration sites.     

Intraguild predation: fiction or reality?

Intraguild predation has become a major research topic in biological control. Quantification of multipredator interactions and an understanding of the consequences on target prey populations are needed, which only highlights the importance of population dynamics models in this field. However, intraguild predation models are usually based on Lotka–Volterra equations, which have been shown not to be adequate for modeling population dynamics of aphidophagous insects and their prey. Here we use a simple model developed for simulation of population dynamics of aphidophagous insects, which is based on the type of egg distribution made by predatory females, to estimate the real strength of intraguild predation in the aphidophagous insects. The model consists of two components: random egg distribution among aphid colonies, and between-season population dynamics of the predatory species. The model is used to estimate the proportion of predatory individuals that face a conflict with a heterospecific competitor at least once during their life. Based on this, predictions are made on the population dynamics of both predatory species. The predictions are confronted with our data on intraguild predation in ladybirds.     

Kin selection and reciprocity in flight formation?

The reasons for conspicuous "V" and other flight formationsin birds are debated. Theory and recent empirical advances showthat energy saving is one important function of flight formations,but some aspects remain poorly understood. Combining theoriesof animal flight and sociality, we suggest that some of thevariation in flight formations has its base in kin selectionand reciprocation. The bird leading an acute V formation savesless energy than does the trailing participants. The disadvantageof leading is reduced in more obtuse formations, and when thelongitudinal distance between neighbors is small, the leadingbird can save about as much energy as others. Therefore, acuteV formations are predicted to occur mainly in circumstancesconducive to kin selection or reciprocity. These mechanismsseem possible, for example, in small flocks of adults with offspring,such as in swans, geese, and cranes. Inclusive fitness advantagesmay then favor an energetically expensive leader role for adults.In small groups, reciprocity is also possible among unrelatedadults that recognize each other and take turns leading theV formation. In contrast, obtuse formations are expected inlarge flocks of unrelated individuals, such as spring flocksof waders migrating long distances. Possibilities for testingthese ideas are discussed.     

Is tadpole coloration adaptive against bird predation?

Birds are visually oriented predators, and some are known or supposed to prey upon tadpoles. Neotropical tadpoles exhibit several colorations that could potentially improve survivorship through camouflage, aposematism, or mimetism. In this study, we tested tadpoles of two hylid species for protective effects of their colorations, one potentially aposematic (Bokermannohyla martinsi) and another (Ololygon machadoi) previously hypothesized as disruptive and/or mimetic to a naucorid insect (Limnocoris porphyros). We conducted in situ and ex situ experiments and observed that the uniformly black B. martinsi tadpoles were more intensely preyed upon by birds than those of O. machadoi, regardless of background color, which may have happened because the disruptive coloration of O. machadoi hampered its detection. Alternatively, the duration of the experiments may have been too short for birds to learn about a possible toxicity of B. martinsi tadpoles, which are more active and thus more visible. Birds that had a previous experience with L. porphyros tended to attack tadpoles of O. machadoi less than naive birds. On the other hand, we did not observe any change in predation levels upon B. martinsi tadpoles whether presented to birds before or after L. porphyros. Our results indicate that the predation by birds, although apparently occasional, is likely to be influenced by tadpole coloration.     

Does copulation increase the risk of predation?

Studies of mating behaviour have assumed that individuals are at greater risk when paired than when engaged in other activities. Recently, four experimental studies of insects and crustaceans have tested this assumption using predators from divergent taxa. Three of these studies indicate that mating carries no additional risk to the participants. Indeed, the findings suggest decreased vulnerability, relative to other activities, due to decreased predation on one or the other of the mating pair.     

Climb and flight speeds of shorebirds embarking on an intercontinental flight; do they achieve the predicted optimal behaviour?

Most Arctic-breeding waders wintering in West Africa cover the first 4000 km of their northward journey in spring by a single flight to western Europe. We examined the extent to which waders economize their flight behaviour during departure by comparing climb rates and forward flight speeds with predictions based on flight mechanic theory and the relevant morphological measurements made of birds collected on the site. With an optical range finder, we followed 98 wader flocks on their departure from Banc d'Arguin in Mauritania. We also measured wind speed and direction at different altitudes by tracking helium-filled balloons and thus were able to deduce airspeeds from groundspeeds of the departing flocks. Of the nine species examined, six showed the predicted negative relationship between climb rate and airspeed, although only one was statistically significant. By normalizing the data, we found a statistically significant negative correlation across all species. Although 17% of the observed climb rates were greater than the predicted theoretical maximum, the average observed climb rate was lower than the predicted optimum and the average observed airspeed was higher. The absolute deviations of climb rates from theory may have been because of the existence of pockets of rising and sinking air at the boundary of desert and ocean. That the absolute deviations in average climb rate and airspeed followed the predicted negative relationship is in accordance with the current theory of flight mechanics.     

Does avian predation risk depress reproduction of voles?

Reproductive output and the growth of captive voles were quantified under high and low avian predation risk in a semi-natural experiment. Voles were exposed to Eurasian kestrels (Falco tinnunculus), the main avian predator of vole species studied (Clethrionomys glareolus, Microtus agrestis and M. rossiaemeridionalis). Vole pairs were housed in cages settled under nest-boxes occupied by breeding kestrels or in control cages settled under empty nest-boxes for 2 weeks. The experiment was conducted in mid-summer when kestrels had half-grown nestlings, because in that time hunting adults and begging nestlings produce noise and scats which may indicate significant predation threat to voles housed underneath the nest-boxes. The risk of kestrel predation did not have any obvious impact on pregnancy rates, mean litter sizes, or growth rates of kestrel-exposed voles compared with control voles studied. These results indicate that the risk of avian predation does not depress the reproductive investment of voles. Received: 3 November 1997 / Accepted: 16 February 1998     

Oddity and the ‘confusion effect’ in predation

We report on two sets of experiments designed to clarify the roles of sensory ‘confusion’ and prey ‘oddity’ as they interact to influence the hunting success of a pursuit predator, the largemouth bass (Micropterus salmoides), on silvery minnows (Hybognathus nuchalis). Bass quickly captured solitary minnows, but performed many unsuccessful attacks and took much longer to make a capture as prey school size was increased. At school sizes of eight and above, bass were effectively stymied, demonstrating the ‘confusion effect’. The inclusion of one or two ‘odd’ (blue-dyed) minnows in a school of eight greatly increased the ability of bass to capture both normal and odd prey, but this effect of oddity disappeared at a school size of 15. The implications of these results for understanding the adaptive basis of mixed species flocks, herds and schools is discussed.     

Is arthropod predation exclusively satiation‐driven?

Functional response models differ in which factors limit predation (e.g. searching efficiency, prey handling time, digestion) and whether predation behaviour is governed by an internal physiological state (e.g. satiation). There is now much evidence that satiation is a key factor in understanding changes in foraging behaviour, and that many predators are effectively digestion limited. Here, we ask if predation in a predatory arthropod can be explained from satiation-driven behaviour alone, or if behaviour is also influenced by the density of prey other than via the effect of prey ingestion on satiation. To address this question a satiation-based predation model is formulated, for which parameters are estimated on the basis of observations on digestion rate, satiation-related prey searching rate and prey capture behaviour, basically under high prey density conditions. The model predictions are subsequently tested against longer term predation experiments carried out at high and low prey densities. Since satiation can easily be linked with egg production, these tests are carried out both for predation and oviposition.
The predator–prey systems under study consist of females of two predatory mite species ( Neoseiulus barkeri and N. cucumeris ) and the larvae of two thrips species ( Thrips tabaci and Frankliniella occidentalis ) as their prey. For N. barkeri foraging on T. tabaci , the model gives good predictions at both high (4 larvae cm⁻¹) and low (0.1–1 larvae cm⁻²) prey densities. For N. cucumeris foraging on F. occidentalis , the predictions hold at the high prey density, but are too low at low prey densities. Thus our analysis indicates that we cannot fully explain density-dependent predation rates from satiation-driven behaviour alone. Different mechanisms are suggested on how prey density may affect foraging efficiency other than via satiation.     

A keystone effect for parasites in intraguild predation?

Intraguild predation (IGP) is common in communities, yet theory suggests it should not often persist and coexistence of participating species should be rare. As parasitism can play keystone roles in interactions between competitors, and between predators and prey, here we examine the role of parasites in maintaining IGP. We used numerical exploration of population dynamic equations to determine coexistence and exclusion zones for two species engaged in IGP with shared parasitism. We demonstrate that parasitism increases the range of conditions leading to coexistence when the parasite exerts a greater deleterious effect on the 'stronger' species in terms of the combined effects of competition and predation. Such a parasite can enable an inferior competitor that is also the less predatory to persist, and may actually lead to numerical dominance of this species.     

Does predation maintain eyespot plasticity in Bicyclus anynana?

The butterfly Bicyclus anynana exhibits phenotypic plasticity involving the wet-season phenotype, which possesses marginal eyespots on the ventral surface of the wings, and the dry-season form, which lacks these eyespots. We examined the adaptive value of phenotypic plasticity of B. anynana in relation to the defence mechanisms of crypsis and deflection. We assessed the visibility differences between spotless and spotted butterflies against backgrounds of brown (dry season) or green (wet season) leaves. Spotless butterflies were highly cryptic and less predated by adult bird predators than were spotted ones when presented against brown leaf litter. However, the advantage of crypsis disappeared in the wet-season habitat as both forms were equally visible. In later experiments, naive birds presented with resting butterflies in the wet-season habitat tended to learn more rapidly to capture spotless butterflies, suggesting a slight selective advantage of possessing eyespots. Moreover, marginal eyespots increased significantly the escape probability of butterflies that were attacked by naive birds compared to those attacked by adult birds, although there were no differences in prey capture success within naive predators. Our results show that natural selection acts against eyespots in the dry season, favouring crypsis, whereas in the wet season it may favour eyespots as deflective patterns.     

Can predation by invasive mice drive seabird extinctions?

The house mouse, Mus musculus, is one of the most widespread and well-studied invasive mammals on islands. It was thought to pose little risk to seabirds, but video evidence from Gough Island, South Atlantic Ocean shows house mice killing chicks of two IUCN-listed seabird species. Mouse-induced mortality in 2004 was a significant cause of extremely poor breeding success for Tristan albatrosses, Diomedea dabbenena (0.27 fledglings/pair), and Atlantic petrels, Pterodroma incerta (0.33). Population models show that these levels of predation are sufficient to cause population decreases. Unlike many other islands, mice are the only introduced mammals on Gough Island. However, restoration programmes to eradicate rats and other introduced mammals from islands are increasing the number of islands where mice are the sole alien mammals. If these mouse populations are released from the ecological effects of predators and competitors, they too may become predatory on seabird chicks.     

Avian predation on a parasitic fly of cervids during winter: can host‐related cues increase the predation risk?

The deer ked (Lipoptena cervi) is an ectoparasitic fly on cervids that has expanded its distribution rapidly in Northern Europe. However, the regulating biotic factors such as predation remain unknown. The host‐independent pupal stage of the fly lasts for several months. Blackish pupae are visible against snow, especially on the bedding sites of hosts, and are thus exposed to predators. To evaluate the role of predation on the invasion dynamics and evolution of L. cervi, we monitored pupal predation on artificial bedding sites in three geographical areas in Finland during winter. We explored: (1) possible predators; (2) magnitude of predation; and (3) whether predation risk is affected by host‐derived cues. We demonstrate that pupae are predated by a number of tit species. Any reddish brown snow discoloration on bedding sites, indicating heavy infestation of the host, serves as an exploitable cue for avian predators, thereby increasing the risk of pupal predation. The ability of tits to use this host‐derived cue seems to be dependent on the prevalence of L. cervi and the period of invasion history, which suggests that it may be a learned behavioural response. Predation by tits may potentially affect the L. cervi population dynamics locally. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 275–286.     

Does land use change influence predation of bird nests?

Worldwide, many areas of agricultural land which were once covered with native vegetation have been converted to tree plantations. Such landscape transformation can influence the dynamics of wildlife populations through, for example, altering rates of predation (e.g. predation of nests of birds). Nest predation can influence reproductive success and, in turn, may alter populations by affecting juvenile recruitment. We quantified predation of bird nests in woodland remnants surrounded by two types of land use, grazing farmland and exotic Radiata pine (Pinus radiata) plantation. We also examined differences in predation rates between artificial and natural nests. We found both artificial and natural nests were more susceptible to nest predation in woodland remnants surrounded by a pine plantation than in woodland remnants located within farmland. Our study suggests that higher levels of nest predation may reduce occupancy of woodland remnants by small‐bodied birds over time, including species of conservation concern. This may have been occurred as a result of the conversion of semi‐cleared grazing land to exotic pine plantation.