Experimental increase of predation risk induces breeding dispersal of Tengmalm's owl

Nest predation and its avoidance are critical components of an individual's fitness and play an important role in life history evolution. Almost all studies on this topic have been observational, and thus have not been able to separate the effects of individual quality, habitat selection and predation risk of given nest sites from each other. More experimental studies on nest predation and breeding dispersal, therefore, are needed to avoid confusing interpretations of the results. In western Finland, pine marten (Martes martes) predation risk was experimentally simulated at the nests of Tengmalm's owls (Aegolius funereus) by using a caged American mink (Mustela vison) as a predator. Nests without exposure to a mink served as controls. In accordance with our predictions and earlier observational studies, males exposed to simulated predation risk increased nest-hole shift and breeding dispersal distances compared to control males. Nest-hole shift and long breeding dispersal distances probably decrease the risk of nest predation, because pine martens are known to revisit nest-holes they have found.     

Variation in digestion of prey by the Tawny owl (Strix aluco)

Two Tawny owl chicks were collected from their nests and observations were made on their food consumption, digestion and pellet production. The female chick escaped after three months in captivity; the male continued to provide data for a further 23 months.     

Tawny owl reproduction and offspring sex ratios under variable food conditions

Tawny owl reproduction and offspring sex ratios have been considered to depend on the abundance of small voles. We studied reproductive performance (laying date, clutch and brood size) during 1995–2003 and offspring sex ratios from 1999 to 2003 in relation to the abundance of small voles and food delivered to the nest in a tawny owl population in southern Finland. Abundance of small voles (field and bank voles) was based on trappings in the field, and estimates of food delivery was based on diet analysis of food remains in the nest boxes. In this population, reproductive output was not related to the abundance of small voles. Analysis of food delivered to the nest showed that the prey weight per offspring varied more than twofold between years and revealed that this difference was mainly related to the proportion of water voles in the diet. Only the number of water voles correlated with laying dates. Offspring sex ratios were weakly male biased (55%) but did not differ from parity. Sex ratios were not related to the abundance of small voles, and we found no evidence that parents delivered more food to nests with proportionally more offspring of the larger (female) sex. Our results underline the notion that populations may differ in their sex allocation pattern, and suggest such differences may be due to diet.     

Factors affecting spontaneous vocal activity of Tawny Owls Strix aluco and implications for surveying large areas

To use vocalizations properly for the estimation of owl population size, it is important to identify how environmental factors affect owl calling behaviour. Here, we analyse how intrinsic and extrinsic factors modify the vocal activity of Tawny Owls Strix aluco in two areas of northern Spain. From March 2013 to May 2015, we radiotracked 20 Tawny Owls and also undertook a systematic survey in which we collected data on spontaneous vocal activity (hoot/call) of the tagged owls, plus their mates and neighbours (36 untagged owls). After 223 nights in Valle de Mena and 224 in Duranguesado we obtained a total of 8791 records of vocal activity. The annual proportion of surveys on which an owl called was 6.3% and did not differ between the study areas. Vocal activity of Tawny Owls varied with sex, annual cycle stage and weather. Male owls were significantly more vocal than females year‐round, and vocal activity peaked during the incubation and post‐breeding periods. Wind and rain adversely affected vocal activity of both sexes throughout the year. Tagged owls were detected more often than untagged owls, which we interpret as an observer effect because the systematic survey ensured that short distances to tagged owls increased the probability of detecting vocal activity. In fact, 2.8% of variation in vocal activity was due to detectability differences between tagged and untagged owls. We conclude that if fieldwork is carried out during the optimum period of the year for vocal detection (i.e. incubation, which in our case was around mid‐April), and under good weather conditions (dry and calm nights), censuses based on spontaneous vocal activity would detect only approximately 12% of the true Tawny Owl population.     

Body weight, gonad development and moult in the Tawny owl (Strix alum)

Gonad development, moult and seasonal changes in body weight and composition in the Tawny owl Strix aluco were studied by examining the carcasses of 369 owls (mostly road casualties) supplemented by 112 weights of live birds. In breeding females laying was preceded by the accumulation of fat and to a lesser extent protein which meant that they weighed more at this time (February/March) than at any other. Females declined in weight after laying but were still heavy during incubation. In contrast, males and non-breeding females did not increase in weight before the start of the breeding season. Juveniles reached or even exceeded adult weight well before independence due to the deposition of fat. Even after the exclusion of diseased or contaminated individuals, 9·4% of the birds examined were identified as starving; most of these were in the autumn and were probably newly-independent young wandering in search of territories. In both sexes gonad maturation was of brief duration coinciding with the period (mid-March to mid-April) in which eggs are normally laid. Ovarian growth was biphasic. In the three months prior to the breeding season ovarian condition in different birds was positively correlated with body weight and it appeared that the largest ovarian follicles of females in poor condition failed to attain the size from which rapid growth to final ovulation occurs. in males testis size in the breeding season was correlated with pectoral muscle weight (an index to protein condition) but not body weight. The majority of adults commenced wing moult in June. The average duration of primary moult was estimated to be 77 days. Healthy birds replaced the primaries of both wings at the same rate but most diseased birds moulted asymmetrically and/or out of season. First-year birds renewed their body feathers between September and November. In the Tawny owl territory establishment, breeding and moult are temporally separated.     

The importance of body reserves for successful reproduction in the Tawny owl (Strix aluco)

One-quarter of Tawny owl nests fail to hatch young, mainly because the eggs are chilled and/or deserted. In 1973–74 automatic photography was employed at four nests near Oxford to relate the incubation behaviour of females to the ration of prey supplied to them by their mates. The eggs did not hatch in two nests and young fledged from only one of the others. Females were less attentive at the nests which failed during incubation and on average received less prey, but even at successful nests there were some nights when the female was supplied with less than her estimated daily food requirement. Female Tawny owls accumulate large reserves of fat and protein before laying. These buffer against any temporary inability of the male to provide sufficient food during incubation, and enable the female to stay on the nest rather than hunt for herself and risk the eggs becoming chilled. When prey are scarce, many females do not lay at all, and the ultimate factor determining whether breeding takes place may be the female's ability to acquire body reserves sufficient to provide a chance of breeding successfully.     

Acoustic and visual adaptations to predation risk: a predator affects communication in vocal female fish

Predation is an important ecological constraint that influences communication in animals. Fish respond to predators by adjusting their visual signaling behavior, but the responses in calling behavior in the presence of a visually detected predator are largely unknown. We hypothesize that fish will reduce visual and acoustic signaling including sound levels and avoid escalating fights in the presence of a predator. To test this we investigated dyadic contests in female croaking gouramis (Trichopsis vittata, Osphronemidae) in the presence and absence of a predator (Astronotus ocellatus, Cichlidae) in an adjoining tank. Agonistic behavior in T. vittata consists of lateral (visual) displays, antiparallel circling, and production of croaking sounds and may escalate to frontal displays. We analyzed the number and duration of lateral display bouts, the number, duration, sound pressure level, and dominant frequency of croaking sounds as well as contest outcomes. The number and duration of lateral displays decreased significantly in predator when compared with no-predator trials. Total number of sounds per contest dropped in parallel but no significant changes were observed in sound characteristics. In the presence of a predator, dyadic contests were decided or terminated during lateral displays and never escalated to frontal displays. The gouramis showed approaching behavior toward the predator between lateral displays. This is the first study supporting the hypothesis that predators reduce visual and acoustic signaling in a vocal fish. Sound properties, in contrast, did not change. Decreased signaling and the lack of escalating contests reduce the fish’s conspicuousness and thus predation threat.     

Maternal transfer of androgens in eggs is affected by food supplementation but not by predation risk

Mothers may affect the future success of their offspring by varying allocation to eggs and embryos. Allocation may be adaptive based on the environmental conditions perceived during early breeding. We investigated the effects of food supplementation and predation risk on yolk hormone transfer in the pied flycatcher Ficedula hypoleuca. In a food supplementation experiment, females were food‐supplemented prior to and during egg‐laying and androgen concentrations were measured throughout the laying order. Predation risk was investigated in three different studies combining both correlative data, where flycatchers bred in close proximity to two different predator species that prey upon adult flycatchers (either Tengmalm's owl Aegolius funereus or pygmy owl Glaucidium passerinum), and an experimental manipulation, where flycatchers were exposed to cues of a nest predator (least weasel Mustela nivalis). Females receiving food supplementation laid eggs with lower concentrations of androstenedione (A4) than females not receiving food supplements. Yolk testosterone (T) concentration showed the same pattern but the difference was not statically significant. Testosterone (but not A4) concentration increased within clutches, from the first to the last egg, independently of the food supplementation. Females breeding under high predation risk did not differ from control females in their yolk androgen levels (A4, T or progesterone). However, concentrations of A4 tended to be lower in the proximity of pygmy owls, which could indirectly increase offspring survival after fledging. Food supplementation during egg‐laying seems to have a stronger impact on maternal transfer of androgens than predation risk. Food availability and predation risk could differentially affect the trade‐offs of androgen allocation for the offspring when raised in good vs. dangerous environments.     

Rodent foraging is affected by indirect, but not by direct, cues of predation risk

We used foraging trays to determine whether oldfield mice, Peromyscuspolionotus, altered foraging in response to direct cues of predationrisk (urine of native and nonnative predators) and indirectcues of predation risk (foraging microhabitat, precipitation,and moon illumination). The proportion of seeds remaining ineach tray (a measure of the giving-up density [GUD]) was usedto measure risk perceived by mice. Mice did not alter theirGUD when presented with cues of native predators (bobcats, Lynxrufus, and red foxes, Vulpes vulpes), recently introduced predators(coyotes, Canis latrans), nonnative predators (ocelots, Leoparduspardalis), a native herbivore (white-tailed deer, Odocoileusvirginianus), or a water control. Rather, GUD was related tomicrohabitat: rodents removed more seeds from foraging trayssheltered beneath vegetative cover compared with exposed traysoutside of cover. Rodents also removed more seeds during nightswith precipitation and when moon illumination was low. Our resultssuggest that P. polionotus used indirect cues rather than directcues to assess risk of vertebrate predation. Indirect cues maybe more reliable than are direct scent cues for estimating riskfrom multiple vertebrate predators that present the most riskin open environments.     

The effects of owl predation on the foraging behavior of heteromyid rodents

Summary Researchers have documented microhabitat partitioning among the heteromyid rodents of the deserts of North America that may result from microhabitat specific predation rates; large/bipedal species predominate in the open/risky microhabitat and small/quadrupedal species predominate in the bush/safer microhabitat. Here, we provide direct experimental evidence on the role of predatory risk in affecting the foraging behavior of three species of heteromyid rodents: Arizona pocket mouse (Perognathus amplus; small/quadrupedal), Bailey's pocket mouse (P. baileyi; large/quadrupedal), and Merriam's kangaroo rat (Dipodomys merriami; large/bipedal). Both kangaroo rats and pocket mice are behaviorally flexible and able to adjust their foraging behavior to nightly changes in predatory risk. Under low levels of perceived predatory risk the kangaroo rat foraged relatively more in the open microhabitat than the two pocket mouse species. In response to the presence of barn owls, however, all three species shifted their habitat use towards the bush microhabitat. In response to direct measures of predatory risk, i.e. the actual presence of owls, all three species reduced foraging and left resource patches at higher giving up densities of seeds. In response to indirect indicators of predatory risk, i.e. illumination, there was a tendency for all three species to reduce foraging. The differences in morphology between pocket mice and kangaroo rats do appear to influence their behavioral responses to predatory risk.     

Snake activity affects seasonal variation in nest predation risk for birds

Variation in predator behavior has been proposed, but not tested, as a mechanism producing seasonal declines in avian nest success. We test this hypothesis by documenting seasonal activity of Texas ratsnakes Elaphe obsoleta and nest failure of endangered black-capped vireos Vireo atricapilla and golden-cheeked warblers Dendroica chrysoparia on which the snakes prey. Nest survival analysis was based on 880 vireo and 228 warbler nests and 3,060 snake locations from 62 radio-tracked snakes. Although nest success varied with snake activity for both birds, specific patterns differed substantially. Vireo daily nest survival was negatively correlated with snake activity over the three-year study, despite substantial variation among years in weather, and the fact that these birds are almost certainly a minor prey species of the ratsnakes. Warblers exhibited less clear-cut seasonal variation in nest success, and the association between nest success and snake activity was less pronounced than for vireos. Increased activity at warmer temperatures explained some of the seasonal change in snake movements, although mating may have accounted for a mid-season peak in activity. These results indicate that variation in predator behavior can be associated with and potentially cause seasonal changes in nest success, but also that these relationships are species specific even within the same community and may depend on aspects of the nesting ecology of the prey such as nest site selection.     

Landscape-dependent response to predation risk by forest birds

Knowing how forest loss and associated fragmentation actually impact individual birds is essential to our understanding of consequences at the population level. We conducted a landscape-level experiment to test whether deforestation affects the trade-off between foraging and antipredatory behaviour of black-capped chickadees ( Poecile atricapilla ) in 24 landscapes (range 8–88% forest cover, 500-m radius) during two winters. At a field-forest edge in the centre of each landscape, we used the maximum distance ventured into the open by flocks to get sunflower seeds placed on the snow-covered fields, as a measure of risk-taking. In the more deforested landscapes, chickadees ventured farther (up to the maximum of 40 m) into the open. Edge density and proportion of conifers in the forest had no influence on risk-taking. However, where ad libitum food was available for a few weeks prior to the experiment (in 12 of the 24 landscapes), chickadees ventured four meters or less away from the forest edge, regardless of the level of deforestation. We conclude that landscape deforestation increases energy stress, which in turn promotes risk-taking, and may therefore increase winter mortality through greater exposure to predators.     

Temporal learning of predation risk by embryonic amphibians

For prey species that rely on learning to recognize their predators, natural selection should favour individuals able to learn as early as possible. The earliest point at which individuals can gather information about the identity of their potential predators is during the embryonic stage. Indeed, recent experiments have demonstrated that amphibians can learn to recognize predators prior to hatching. Here, we conditioned woodfrog embryos to recognize predatory salamander cues either in the morning or in the evening, and subsequently exposed the two-week-old tadpoles to salamander cues either in the morning or in the evening, and recorded the intensity of their antipredator behaviour. The data indicate that amphibians learn to recognize potential predators while still in the egg, and also learn the temporal component of this information, which they use later in life, to adjust the intensity of their antipredator responses throughout the day.     

Tawny owl prey remains indicate differences in the dynamics of coastal and inland vole populations in southern Finland

Some studies suggest that mild winters decrease overwinter survival of small mammals or coincide with decreased cyclicity in vole numbers, whereas other studies suggest non-significant or positive relationships between mild winter conditions and vole population dynamics. We expect for the number of voles to be higher in the rich and low-lying habitats of the coastal areas than in the less fertile areas inland. We assume that this geographical difference in vole abundances is diminished by mild winters especially in low-lying habitats. We examine these relationships by generalized linear mixed models using prey remains of breeding tawny owls Strix aluco as a proxy for the abundance of voles. The higher number of small voles in the coastal area than in the inland area suggest that vole populations were denser in the coastal area. Vole populations of both areas were affected by winter weather conditions particularly in March, but these relationships differed between areas. The mild ends of winter with frequent fluctuations of the ambient temperature around the freezing point (“frost seesaw”) constrained significantly the coastal vole populations, while deep snow cover, in general after hard winters, was followed by significantly lowered number of voles only in the inland populations. Our results suggest that coastal vole populations are more vulnerable to mild winters than inland ones. We also show that tawny owl prey remains can be used in a meaningful way to study vole population dynamics.     

Human activities change marine ecosystems by altering predation risk

In ocean ecosystems, many of the changes in predation risk – both increases and decreases – are human‐induced. These changes are occurring at scales ranging from global to local and across variable temporal scales. Indirect, risk‐based effects of human activity are known to be important in structuring some terrestrial ecosystems, but these impacts have largely been neglected in oceans. Here, we synthesize existing literature and data to explore multiple lines of evidence that collectively suggest diverse human activities are changing marine ecosystems, including carbon storage capacity, in myriad ways by altering predation risk. We provide novel, compelling evidence that at least one key human activity, overfishing, can lead to distinct, cascading risk effects in natural ecosystems whose magnitude exceeds that of presumed lethal effects and may account for previously unexplained findings. We further discuss the conservation implications of human‐caused indirect risk effects. Finally, we provide a predictive framework for when human alterations of risk in oceans should lead to cascading effects and outline a prospectus for future research. Given the speed and extent with which human activities are altering marine risk landscapes, it is crucial that conservation and management policy considers the indirect effects of these activities in order to increase the likelihood of success and avoid unfortunate surprises.