Helminth Communities of Owls (Strigiformes) Indicate Strong Biological and Ecological Differences from Birds of Prey (Accipitriformes and Falconiformes) in Southern Italy

We compared the helminth communities of 5 owl species from Calabria (Italy) and evaluated the effect of phylogenetic and ecological factors on community structure. Two host taxonomic scales were considered, i.e., owl species, and owls vs. birds of prey. The latter scale was dealt with by comparing the data here obtained with that of birds of prey from the same locality and with those published previously on owls and birds of prey from Galicia (Spain). A total of 19 helminth taxa were found in owls from Calabria. Statistical comparison showed only marginal differences between scops owls (Otus scops) and little owls (Athene noctua) and tawny owls (Strix aluco). It would indicate that all owl species are exposed to a common pool of ‘owl generalist’ helminth taxa, with quantitative differences being determined by differences in diet within a range of prey relatively narrow. In contrast, birds of prey from the same region exhibited strong differences because they feed on different and wider spectra of prey. In Calabria, owls can be separated as a whole from birds of prey with regard to the structure of their helminth communities while in Galicia helminths of owls represent a subset of those of birds of prey. This difference is related to the occurrence in Calabria, but not Galicia, of a pool of ‘owl specialist’ species. The wide geographical occurrence of these taxa suggest that local conditions may determine fundamental differences in the composition of local communities. Finally, in both Calabria and Galicia, helminth communities from owls were species-poor compared to those from sympatric birds of prey. However, birds of prey appear to share a greater pool of specific helmith taxa derived from cospeciation processes, and a greater potential exchange of parasites between them than with owls because of phylogenetic closeness.     

Bird predation by tawny owls (Strix aluco L.) and its effect on the reproductive performance of tits

The density of great tit Parus major L. and blue tit Parus caeruleus L. was artificially increased by placing nest-box colonies for these species in the vicinity of the nests of breeding tawny owls during 1993–1997. Bird prey composition in the owl nests, the proportion of parents disappearing from the breeding tit populations and the reproductive performance of the widowed parents were analysed. The frequency of predation on tits by tawny owls was greater in areas where tit density had been artificially increased. Owls preyed more on tits during the feeding period of owlets than during the incubation period and more in years when snow covered the ground during the incubation period than when it did not. Mortality due to predation was male biased and more females lost their mates in populations breeding near tawny owl nests. Reproductive performance of the widowed parents was lower and their body weights were lighter at the end of the nestling period than those found in birds rearing youngs with their mates. Predation by owls increased the between-year turnover in the breeding tit population: widowed parents did not return to the nesting site for the next breeding season.     

Tawny owl vocal activity is constrained by predation risk

The vocal behaviour of birds may be influenced by many factors, including the risk of being detected by a predator. In Doñana Protected Area, the tawny owl co‐exists alongside its intraguild predator, the eagle owl Bubo bubo. We considered four scenarios to study the vocal behaviour of tawny owls at dusk by analysing: A) the calling rate of all males in 29 sites; B) the calling rate at dusk of males living within the home range of the intraguild predator; C) the calling rate of males living within the home range of the intraguild predator between 60 and 90 min after sunset; and D) the duration of male vocal bouts in visits where eagle owls have called. In scenario A we found that only the number of conspecific males affected the calling rate of tawny owls. In scenario B we observed that the presence of an eagle owl calling constrained the calling rate of the intraguild prey. In scenario C we found that this effect seemed mostly associated to a contemporaneous detection of the intraguild predator’s calls. Finally, in scenario D we found no significant effects on bout duration. These results seem to indicate that tawny owls use their intraguild predator’s calls as a cue to assess predation risk, and then adjust their vocal behaviour in order to minimize predation risk by a predator that may locate its prey by its vocalizations.     

<Emphasis Type="Italic">Apodemus</Emphasis> mice as the main prey that determines reproductive output of tawny owl (<Emphasis Type="Italic">Strix aluco</Emphasis>) in Central Europe

During the years 2008–2014, we studied diet composition, the number of breeding pairs, and reproductive output of tawny owls in Central Europe (Czech Republic) in relation to availability of main prey in the field. We also performed a meta-analysis on diet composition of tawny owl in Europe that confirmed the important role of Apodemus mice in tawny owl diet in Central Europe. In concordance, Apodemus mice were the main prey of tawny owl in our study area (38.7%), and Microtus/Myodes voles (15.4%), birds (12.1%) and others (33.8%) were alternative prey. We found a positive relationship between the proportion of Apodemus mice in the diet and their abundance in the field (beta = 0.23, P = 0.001). Availability of main prey (Apodemus mice, Microtus/Myodes voles or Sorex shrews) in the field was not correlated with the number of breeding pairs. Proportion of birds in diet (expressed by scores from multivariate analysis), which was inversely related to proportion of Apodemus mice, was positively correlated with laying date (beta = 0.66, P = 0.012) and negatively correlated with clutch size (beta = − 0.45, P = 0.004) and brood size (beta = − 0.16, P = 0.076). We also found negative relationships between laying date and clutch size (beta = − 0.13, P = 0.014) and brood size (beta = − 0.07, P = 0.057). Our results support the idea that diet and breeding ecology of owls in Central Europe is mainly driven by the availability of Apodemus mice that are suitable prey due to their similar habitat requirements and nocturnal activity.     

Long-term changes in abundance of bats as revealed by their frequency in tawny owls’ diet

Percentage of bats in tawny owls’ diet was compared in three periods: I — before 1976, II — 1976–1992, III — 1993–2009, by using the published and unpublished material from Poland (only samples over 100 vertebrate prey items). This species of owl showed an opportunistic predation on bats and took them more frequently in periods of higher abundance. Before the mass use of toxic pesticides in Poland, in the period I bats constituted more than 2% of vertebrates in four out of five diet samples (median 2.4%). The lowest bat abundance occurred in Poland in the 1980s and resulted in the lowest percentage of bats taken by owls in the period II (n = 11, median 0.2%). Due to the recovery of bat populations in the period III, the percentage of bats in tawny owls’ diet increased (n = 23, median 0.7%). In large samples (over 200 vertebrate items, n = 21) collected in central and north-eastern Poland the percentage of bats increased from 1980 to 2009 (the estimated average value at the end of that period slightly exceeded 1%). Samples collected at the same five sites in 1975–1992 and again in 2000–2009, confirmed the increasing trend in percentage of bats captured by tawny owls noted in last years.     

Interspecific offspring killing in owls

In Baranya County (Southern Hungary), tawny owls (Strix aluco) and barn owls (Tyto alba) sequentially use the same nest boxes in a significant number of cases. A total of 460 broods were observed between 1996 and 2003 and, in 12 cases, whole broods of dead tawny owl chicks were registered that had apparently been killed. On investigating the reproductive life characteristics, population sizes, and frequency of killing in these two species, it was concluded that: (1) with growing barn owl population, the number of sequential broods increases but changes in tawny owl population size have no effect on the frequency of sequential broods; (2) the number of killings depends on the number of sequential broods; and (3) with growing barn owl population, the number of killings also grows and this change is unaffected by the size of the tawny owl population. However, no killing occurs as long as 50–60% of the nest boxes are unoccupied. There is no killing either until the percentage of nest boxes occupied by barn owls reaches 40%, although a threshold value like this cannot be shown for the tawny owl. In the cases when a barn owl breeding follows the tawny owl's, the percentage of killing is significantly higher compared to that when barn owls do not breed in the same box. These results indicate that barn owls kill the offspring of tawny owls. By these means, they obtain a breeding place earlier than without killing the chicks of the other species, and this results in higher reproductive success. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 488–494.     

Indirect Costs of Parasitism are Shaped by Variation in the Type of Immune Challenge and Food Availability

Parasites can inflict indirect fitness costs to their hosts by eliciting costly immune responses. These costs depend on the type and amount of immunostimulants presented to the host immune system but also on the amount of resources available to fuel host immune responses. Here, we investigated how the relative costs of two different types of immune challenge are modulated by variation in food availability. We injected nestling tawny owls (Strix aluco) with either 10 μg of phytohaemagglutinin (PHA) or 20 μg of lipopolysaccharide (LPS), and subsequently raised them under two different food regimes (food-restricted vs. ad libitum). After controlling for food consumption, we found that LPS-injected nestlings lost more body mass than PHA-injected ones only when food-restricted. We also found that body mass gain of owlets fed ad libitum decreased with the intensity of the skin swelling response against LPS, but not PHA. These experimental and correlative results suggest that nestling tawny owls suffered greater immune costs when treated with LPS than PHA, and that variation in the costs of two different types of immune challenge can be exacerbated under conditions of low food availability. Our study highlights the importance of taking into consideration the interplay between host immunity and nutrition in the study of indirect costs of parasitism.     

Does feeding effort of Tengmalm's owls reflect offspring survival prospects in cyclic food conditions?

In an environment fluctuating in a predicatable manner with wide among-year variation in offspring mortality, fitness is largely influenced by the timing of reproductive investment. In vole-eating nocturnal Tengmalm's owls (Aegolius funereus), within-cycle variation in 1st-year survival of owlets is 3-fold as estimated by the recruitment probability of an offspring. It increases from the peak through the low to the increase phase of the vole cycle. We recorded prey delivery rates of males during a 3-year vole cycle using 4 h continuous recording at night. Males brought significantly more prey items per offspring in a low-vole year than in the increase and peak vole years. In the early night (10 p.m.–12 p.m.), males fed their families equally in the increase and peak vole years, whereas in the late night (0.01–2.00 a.m.) males reduced their feeding rate in the peak year but not in other years. Both prey number and prey mass per offspring were larger in the low and increase vole years than in the peak year, though in the peak phase food is most abundant. We suggest that feeding effort of site-tenacious, long-lived (mean lifespan 3.5 years) male owls culminates in the increase rather than in the peak phase of the vole cycle, because offspring survive better in the former phase.     

Does variation of sex ratio enhance reproductive success of offspring in tawny owls (Strix aluco)

Tawny owls, Strix aluco, laid female-biased clutches on territories with more abundant prey (field voles) in June, the month that chicks fledge. This appeared to enhance the subsequent reproductive success of fledglings, as in 1995 there was a significant correlation between the number of chicks fledged by adult females and the June vole abundance in the territory on which they were reared as chicks. This relationship did not hold for males. Since tawny owls lay eggs in March, these results indicate that owls are able to predict the June vole numbers on their territory, and respond by producing more of the sex most likely to gain a long-term benefit when resources are good.     

Seasonal and habitat variation in the diet of the tawny owl ( Strix aluco ) in central Poland during unusually warm years

In the temperate climate zone in Europe the composition of the diet of predatory vertebrates shows evident variability between the warm and cold season. However, the recently observed climate warming can mitigate the effect of snow cover and low temperatures on the winter foraging ecology of raptors, thus affecting trophic webs in ecosystems. We analysed diet variability in the tawny owl Strix aluco, between the warm and cold seasons of four unusually warm years (as compared to reference years of 1950–2000) in two habitats (forest vs. farmland) in Central Poland. The most important prey group in the tawny owl’s diet were mammals, constituting over 80% of prey items. There were distinct diet differences between the two seasons: insectivorous mammals, birds and amphibians were caught more often during the warm season, and Muridae and Arvicolidae during the cold season. The proportion of insectivorous mammals, voles and amphibians was significantly higher in forest than in farmland. Diet diversity, analysed with rarefaction methods and expressed as the expected cumulative mammal species number for a given number of randomly sampled preyed mammals, was independent of season and higher in forest than in the agricultural habitat. We conclude that even during unusually warm years tawny owls change significantly their feeding habits between the warm and cold season. The effect of season, habitat and weather factors on diet variability in raptors are discussed.     

Perfect is best: low leaf fluctuating asymmetry reduces herbivory by leaf miners

Fluctuating asymmetry (FA) represents small, random variation from symmetry and can be used as an indicator of plant susceptibility to herbivory. We investigated the effects of FA of two oak species, Quercus laevis and Q. geminata, and the responses of three herbivore guilds: leaf miners, gallers, and chewers. To examine differences in FA and herbivory between individuals, 40 leaves from each tree were collected, and FA indices were calculated. To examine differences in FA and herbivory within-individuals, we sampled pairs of mined and unmined leaves for asymmetry measurements. Differences in growth of leaf miners between leaf types were determined by tracing 50 mines of each species on symmetric leaves and asymmetric leaves. Asymmetric leaves contained significantly lower concentrations of tannins and higher concentrations of nitrogen than symmetric leaves for both plant species. Both frequency of asymmetric leaves on plants and levels of asymmetry positively influenced the abundance of Brachys, Stilbosis and other leaf miners, but no significant relationship between asymmetry and herbivory was observed for Acrocercops. Brachys and Stilbosis mines were smaller on asymmetric leaves, but differences in mine survivorship between symmetric and asymmetric leaves were observed only for Stilbosis mines. This study indicated that leaf miners might use leaf FA as a cue to plant quality, although differential survivorship among leaf types was not observed for all species studied. Reasons for the different results between guilds are discussed.     

Analysis of genetic parentage in the tawny owl ( Strix aluco ) reveals extra-pair paternity is low

We have investigated genetic parentage in a Swiss population of tawny owls (Strix aluco). To this end, we performed genetic analysis for six polymorphic loci of 49 avian microsatellite loci tested for cross-species amplification. We found one extra-pair young out of 137 (0.7%) nestlings in 37 families (2.7%). There was no intra-specific brood parasitism. Our results are in accordance with previous findings for other raptors and owls that genetic monogamy is the rule. Female tawny owls cannot raise offspring without a substantial contribution by their mates. Hence one favoured hypothesis is that high paternal investment in reproduction selects for behaviour that prevents cuckoldry. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Host use and developmental instability in the cactophilic sibling species Drosophila gouveai and D. antonietae

The Drosophila repleta group encompasses an ensemble of species that inhabit desertic areas that are inhospitable to other drosophilids. These species have a tractable ecology, as they breed and feed on necrotic tissues of a wide diversity of species of Cactaceae, with a certain degree of host specificity, which makes them suitable models to investigate the role of host plant shifts in diversification. Most species have their own primary host plant, which may be shared with a closely related species. However, the consequences of host plant shifts from primary to secondary hosts have not been thoroughly studied so far. We investigated the effects of the cactus host on developmental instability and performance of D. gouveai Tidon‐Sklorz & Sene and D. antonietae Tidon‐Sklorz & Sene (Diptera: Drosophilidae), a pair of closely related sibling species, and of their F1 interspecific hybrids reared in primary and secondary host plants. Fluctuating asymmetry (FA) for wing size was significantly greater in flies of both species reared in their respective secondary cactus host than in those grown in the primary host. Interspecific hybrids also exhibited host‐dependent levels of FA. However, hybrids did not show greater FA than the parental species, suggesting that hybridization per se did not affect developmental stability. Even though cactus host shifts affected different measures of fitness (larval viability and adult size) both in D. antonietae and D. gouveai, we did not detect a consistent relationship between FA and fitness. Our results suggest that FA cannot be used as a sensitive indicator of genetic and phenotypic quality and we discuss its application as a fitness predictor.     

No relationship between fluctuating asymmetry and fitness in Lychnis viscaria

Several earlier studies have indicated a negative relationship between fluctuating asymmetry (FA) and fitness. We tested this assumption by investigating the association between petal asymmetry and several fitness-related characters among natural and common garden populations of Lychnis viscaria. Neither seed set, germination percentage nor the growth rate of seedlings were related to the level of flower asymmetry either among natural populations or in common garden conditions. The only significant association found was a positive connection between petal asymmetry and seed mass measured from natural populations. Thus, in contrary to many earlier published reports, we did not find any evidence for a negative relationship between FA and fitness even if we controlled for measurement error, we had adequate sample size and we measured these characters in two environments. This suggests that FA is not consistently related to individual quality and fitness.     

Activity patterns at the Arctic Circle: nocturnal eagle owls and interspecific interactions during continuous midsummer daylight

Circadian rhythms result from adaptations to biotic and abiotic environmental conditions that cycle through the day, such as light, temperature, or temporal overlap between interacting species. At high latitudes, close to or beyond the polar circles, uninterrupted midsummer daylight may pose a challenge to the circadian rhythms of otherwise nocturnal species, such as eagle owls Bubo bubo. By non‐invasive field methods, we studied eagle owl activity in light of their interactions with their main prey the water vole Arvicola amphibius, and their competitor the white‐tailed eagle Haliaeetus albicilla during continuous midsummer daylight on open, treeless islands in coastal northern Norway. We evaluated circadian rhythms, temporal overlap, exposure, and spatial distribution. The owls maintained a nocturnal activity pattern, possibly because slightly dimmer light around midnight offered favourable hunting conditions. The eagles were active throughout the 24‐h period as opposed to the strictly diurnal rhythm reported elsewhere, thus increasing temporal overlap and the potential for interference competition between the two avian predators. This may indicate an asymmetry, with the owls facing the highest cost of interference competition. The presence of eagles combined with constant daylight in this open landscape may make the owls vulnerable to interspecific aggression, and contrary to the available literature, eagle owls rarely exposed themselves visually during territorial calls, possibly to avoid detection by eagles. We found indications of spatial segregation between owls and eagles reflecting differences in main prey, possibly in combination with habitat‐mediated avoidance. Eagle owl vocal activity peaked in the evening before a nocturnal peak in visual observations, when owls were active hunting, consistent with the hypothesis of a dusk chorus in nocturnal bird species. The owls may have had to trade‐off between calling and foraging during the few hours around midnight when slightly dimmer light reduced the detection risk while also providing better hunting conditions     

Habitat dependent associations between parasitism and fluctuating asymmetry among endemic stickleback populations

Inconsistencies in the relationship between fluctuating asymmetry (FA) and fitness may be due to selection acting on the degree of trait asymmetry that differs among populations or among traits. We assessed relationships between parasite susceptibility and fluctuating asymmetry in the number of bony lateral plates among 83 populations of freshwater Gasterosteus aculeatus (three spined stickleback) and among lateral plate positions that vary in the selection they experience for symmetry. The correlation between FA and parasite infection was highly variable among samples. Excess of infected asymmetric G. aculeatus increased significantly as the robustness of structural predator defences decreased. This effect was found for one parasite species only (Eustrongylides sp.) and was slightly stronger in females. In addition, there was a trend for there to be an excess of infected females asymmetric in those lateral plates positions that did not experience selection for their symmetry, although the trend only approached significance. These results suggest that selection for trait symmetry can obscure relationships between fitness and individual-wide developmental stability, providing one possible explanation for some of the heterogeneity in FA/fitness relationships seen in the literature. These results are also consistent with previous reports showing that ecological segregation between symmetric and asymmetric G. aculeatus and between sexes can alter the FA/fitness relationship.     

Strong nonlinear selection against fluctuating asymmetry in wild populations of a marine fish

Theoretical links between fluctuating asymmetry (FA) and fitness have led many to use FA as a proxy for average fitness. However, studies examining whether asymmetry actually correlates with individual fitness in wild populations are relatively rare and often use simple measures of association (e.g., correlation coefficients). Consequently, the pattern of selection on asymmetry in the wild is seldom clear. We examined selection on FA of pectoral fin morphology in two wild populations of a marine fish (the kelp perch; Brachyistius frenatus). As expected, variance in signed FA in each initial sample was significantly greater than that found in the surviving population, indicating selection against FA. Our estimate of the fitness surface confirmed perfect symmetry as the phenotypic optimum and indicated strong, nonlinear selection against asymmetry. No difference in the form of selection was detected between populations. However, the level of FA in the initial samples varied among populations, leading to an overall difference in the level of selective mortality. Our results suggest that selection on asymmetry in wild populations may be strongly nonlinear, and indicate that the demographic costs of asymmetry may play a substantial role in the dynamics of populations.     

Increased autumn rainfall disrupts predator–prey interactions in fragmented boreal forests

There is a pressing need to understand how changing climate interacts with land‐use change to affect predator–prey interactions in fragmented landscapes. This is particularly true in boreal ecosystems facing fast climate change and intensification in forestry practices. Here, we investigated the relative influence of autumn climate and habitat quality on the food‐storing behaviour of a generalist predator, the pygmy owl, using a unique data set of 15 850 prey items recorded in western Finland over 12 years. Our results highlighted strong effects of autumn climate (number of days with rainfall and with temperature <0 °C) on food‐store composition. Increasing frequency of days with precipitation in autumn triggered a decrease in (i) total prey biomass stored, (ii) the number of bank voles (main prey) stored, and (iii) the scaled mass index of pygmy owls. Increasing proportions of old spruce forests strengthened the functional response of owls to variations in vole abundance and were more prone to switch from main prey to alternative prey (passerine birds) depending on local climate conditions. High‐quality habitat may allow pygmy owls to buffer negative effects of inclement weather and cyclic variation in vole abundance. Additionally, our results evidenced sex‐specific trends in body condition, as the scaled mass index of smaller males increased while the scaled mass index of larger females decreased over the study period, probably due to sex‐specific foraging strategies and energy requirements. Long‐term temporal stability in local vole abundance refutes the hypothesis of climate‐driven change in vole abundance and suggests that rainier autumns could reduce the vulnerability of small mammals to predation by pygmy owls. As small rodents are key prey species for many predators in northern ecosystems, our findings raise concern about the impact of global change on boreal food webs through changes in main prey vulnerability.     

Natal conditions alter age-specific reproduction but not survival or senescence in a long-lived bird of prey

1. Natal conditions and senescence are two major factors shaping life-history traits of wild animals. However, such factors have rarely been investigated together, and it remains largely unknown whether they interact to affect age-specific performance. 2. We used 27 years of longitudinal data collected on tawny owls with estimates of prey density (field voles) from Kielder Forest (UK) to investigate how prey density at birth affects ageing patterns in reproduction and survival. 3. Natal conditions experienced by tawny owls, measured in terms of vole density, dramatically varied among cohorts and explained 87% of the deviance in first-year apparent survival (annual estimates ranging from 0·07 to 0·33). 4. We found evidence for senescence in survival for females as well as for males. Model-averaged estimates showed that adult survival probability declined linearly with age for females from age 1. In contrast, male survival probability, lower on average than for female, declined after a plateau at age 1-3. 5. We also found evidence for reproductive senescence (number of offspring). For females, reproductive performance increased until age 9 then declined. Males showed an earlier decline in reproductive performance with an onset of senescence at age 3. 6. Long-lasting effects of natal environmental conditions were sex specific. Female reproductive performance was substantially related to natal conditions (difference of 0·24 fledgling per breeding event between females born in the first or third quartile of vole density), whereas male performance was not. We found no evidence for tawny owls born in years with low prey density having accelerated rates of senescence. 7. Our results, combined with previous findings, suggest the way natal environmental conditions affect senescence varies not only across species but also within species according to gender and the demographic trait considered.     

Parent-offspring conflict over duration of parental care and its consequences in tawny owls Strix aluco

In species defending territories, fitness of newly independent juveniles could depend on phenotypic quality or early access to vacant resources as a result of rapid dispersal. In the first case, parent-offspring conflict will arise when parents cease feeding maturing offspring while these still demand provisioning. In the latter case, the young should decide when to stop begging to search for vacant territories. I radio-tracked 72 juvenile tawny owls from independence to sexual maturity to investigate whether parents or offspring decided the timing of the onset of independence and compare effects of age versus date of independence on survival and reproductive status. Juvenile owls stopped begging when 90–123 d old. This was synchronous within broods, independent of gender or age rank. Independence age of cross-fostered young varied across foster nests but was independent of hatching nest. After independence, young roosted in their parents' territory for 18 d on average before dispersal. This suggests that juvenile owls would rather extend dependency than disperse early. As predicted from this behaviour, age of independence had a positive influence on survival and reproductive status at maturity, whereas there were no effects of date of independence.