The role of Apodemus mice and Microtus voles in the diet of the Tengmalm’s owl in Central Europe

Based on a long-term dataset (1999–2010), we investigated how the availability of main prey affects the breeding density and food ecology of the Tengmalm’s owl (Aegolius funereus) in the Czech Republic. In particular, we assessed the role of Microtus voles and Apodemus mice in the diet, based on the main predictions of the optimal diet theory that the diet composition depends on the availability of the main prey. We found that (i) the Tengmalm’s owl exhibited no numerical response to the availability of Microtus voles and Apodemus mice in the field; (ii) the availability of Apodemus mice in the field positively affected their proportion in the diet (26 %), and despite a high proportion of Microtus voles in the owls’ diet (47 %), no relationship was found between their availability in the field and proportion in the diet; (iii) the proportion of Apodemus mice was negatively correlated to the proportion of Microtus voles, Sorex shrews and birds in the diet, but no similar relationship was detected for Microtus voles; (iv) the reproductive output of Tengmalm’s owls was positively correlated to the proportion of Apodemus mice in the diet, as well as to Apodemus mice and Microtus vole availability in the field; and (v) diet diversity and diet overlap were not significantly affected by the abundance of Apodemus mice and Microtus voles. Therefore, the validity of these main optimal diet theory predictions was not confirmed, especially for Microtus vole prey, due to an opportunistic choice between Apodemus mice and Microtus voles. We suggest that the reproductive output of nocturnal raptors in Central Europe may be less dependent on Microtus vole supply than that of their northern counterparts.     

<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.     

The relationship between intra–guild diet overlap and breeding in owls in Israel

Even though intra-guild predators frequently prey on the same species, it is unclear whether diet overlap between two predators is a source of interspecific competition or whether predators simply use the same abundant prey resource. We measured the extent to which the diets of barn owls (Tyto alba) and long-eared owls (Asio otus) in Israel overlap and examined whether yearly differences in diet overlap correlate with barn owl breeding success. Pianka’s index of niche overlap was positively related to barn owl population size but not to its breeding success. The number of breeding barn owls was higher when long-eared owls consumed more rodents, suggesting that diet overlap most likely increased when rodents became more abundant. Therefore, in Israel, when these two owl species prey more often on rodents, their diets are more similar and interspecific competition is reduced. Unlike sympatric populations in Europe, in years when rodents are less abundant in Israel long-eared owls switch to hunting alternative prey (e.g., birds), perhaps to avoid competition with barn owls.     

Do Tengmalm’s Owls alter parental feeding effort under varying conditions of main prey availability?

We studied the diet composition and behavioural responses to variable food conditions in Tengmalm’s Owls (Aegolius funereus). The abundance of main prey (voles and mice) of owls was higher in the Ore Mountains, Czech Republic, than in the Kauhava region, Finland. We monitored nests continuously by a camera system to estimate the feeding frequency and to identify prey items provided to nestlings. We recorded 990 prey deliveries at six nests in the Ore Mountains and 1,679 prey deliveries at nine nests in the Kauhava region. Mice (Apodemus) and voles (Microtus and Clethrionomys) were the main foods of owls in the Ore Mountains, whereas voles (Clethrionomys and Microtus) and shrews (Sorex) were the main foods in the Kauhava region. In consequence, on average smaller prey items were brought to nestlings at the Finnish site. However, both absolute and relative (per one nestling) feeding frequency was higher in the Kauhava region, and the biomass available to individual nestlings did not differ between the two areas. Moreover, the Finnish and Czech pairs produced about the same number of fledglings. Our results suggest that male owls are able to maintain the amount of food required for chicks by switching to alternative prey, and to increase their prey delivery rates under conditions of reduced abundance of main food.     

Variation in the diet composition of a generalist predator, the red fox, in relation to season and density of main prey

Diet composition of a generalist predator, the red fox (Vulpes vulpes) in relation to season (winter or summer) and abundance of multi-annually cyclic voles was studied in western Finland from 1983 to 1995. The proportion of scats (PS; a total of 58 scats) including each food category was calculated for each prey group. Microtus voles (the field vole M. agrestis and the sibling vole M. rossiaemeridionalis) were the main prey group of foxes (PS = 0.55) and they frequently occurred in the scats both in the winter and summer (PSs 0.50 and 0.62, respectively). There was a positive correlation between the PSs of Microtus voles in the winter diet of foxes and the density indices of these voles in the previous autumn. Other microtine rodents (the bank vole Clethrionomys glareolus, the water vole Arvicola terrestris and the muskrat Ondatra zibethicus) were consumed more in winter than in summer. The unusually high small mustelid predation by red foxes (PS = approx. 0.10) in our study area gives qualitative support for the hypothesis on the limiting impact of mammalian predators on least weasel and stoat populations. None of the important prey groups was preyed upon more at low than at high densities of main prey (Microtus voles). This is consistent with the notion that red foxes are generalist predators that tend to opportunistically subsist on many prey groups. Among these prey groups, particularly hares and birds (including grouse), were frequently used as food by foxes.     

Responses of stoats and least weasels to fluctuating food abundances: is the low phase of the vole cycle due to mustelid predation?

Summary We studied responses of stoats and least weasels to fluctuating vole abundances during seven winters in western Finland. Density indices of mustelids were derived from snow-tracking, diet composition from scat samples, and vole abundances from snap-trapping. Predation rate was estimated by the ratio of voles to mustelids and by the vole kill rate by predators (density of predator x percentage of voles in the diet). We tested the following four predictions of the hypothesis that small mustelids cause the low phase of the microtine cycle. (1) The densities of predators should lag well behind the prey abundances, as time lags tend to have destabilizing effects. The densities of stoats fluctuated in accordance with the vole abundances, whereas the spring densities of least weasels tracked the vole abundances with a half-year lag and the autumn densities with a 1-year lag. (2) Predators should not shift to alternative prey with declining vole densities. The yearly proportion of Microtus voles (the staple prey) in the diet of stoats varied widely (range 16–82%) and was positively correlated with the winter abundance of these voles. In contrast, the same proportion in the food of least weasels was independent of the vole abundance. (3) The ratio of voles to small mustelids should be smallest in poor vole years and largest in good ones. This was also observed. (4) Vole densities from autumn to spring should decrease more in those winters when vole kill rates are high than when they are low. The data on least weasels agreed with this prediction. Our results from least weasels were consistent with the predictions of the hypothesis, but stoats behaved like semi-generalist predators. Accordingly, declines and lows in the microtine cycle may be due to least weasel predation, but other extrinsic factors may also contribute to crashes.     

The effects of sex,age, season and habitat on diet of the red fox <Emphasis Type="Italic">Vulpes vulpes</Emphasis> in northeastern Poland

The diet of the red fox Vulpes vulpes was investigated in five regions of northeastern Poland by stomach content analysis of 224 foxes collected from hunters. The red fox is expected to show the opportunistic feeding habits. Our study showed that foxes preyed mainly on wild prey, with strong domination of Microtus rodents, regardless of sex, age, month and habitat. Voles Microtus spp. were found in 73% of stomachs and constituted 47% of food volume consumed. Other food items were ungulate carrion (27% of volume), other mammals (11%), birds (9%), and plant material (4%). Sex- and age-specific differences in dietary diversity were found. Adult males and juvenile foxes had larger food niche breadths than adult females and their diets highly overlapped. Proportion of Microtus voles increased from autumn to late winter. Significant habitat differences between studied regions were found. There was a tendency among foxes to decrease consumption of voles with increasing percentage of forest cover. Based on our findings, red foxes in northeastern Poland can be recognized as a generalist predators, consuming easily accessible and abundant prey. However, high percentage of voles consumed regardless of age, sex, month, or habitats may indicate red fox specialization in preying on Microtus rodents.     

Food habits of stoatsMustela erminea and weaselsMustela nivalis in Denmark

StoatsMustela erminea Linnaeus, 1758 and weaselsMustela nivalis Linnaeus, 1766 exploit the same array of prey species at different frequencies according to body size. The rabbitOryctolagus cuniculus, which typically is the dominant prey for stoats in temperate parts of Europe, is absent in Denmark. The present study based on gastrointestinal tract contents examines the food habits of sympatric stoats (n=112) and weasels (n=132) from Denmark. Rodents were the most important prey group for both stoats and weasels, constituting 77% and 84% of their diet respectively, expressed as frequency of occurrence. No differences were detected in the proportions of major prey groups between sexes within species, but between species the diets differed. Stoats ate birds and birds’ eggs more often than weasels did, while weasels ate more insectivores. Stoats ate more oftenMicrotus voles and water voleArvicola terrestris than weasels did, while weasels ate more bank voleClethrionomys glareolus and moleTalpa europaea. There was a larger dietary overlap between sympatric stoats and weasels in Denmark than in other dietary studies in areas where rabbits were available.     

The kestrel (Falco tinnunculus L.) in Berlin: investigation of breeding biology and feeding ecology

Approximately 200–250 pairs of kestrels (Falco tinnunculus) breed in Berlin, preferentially in nest boxes. From 2002 to 2004, ten monitoring sites (breeding sites) characterised by different housing structure, land utilisation, vegetation cover and degree of building density were studied in Berlin: four in the city centre, three in a mixed zone and three in the outskirts. All pairs bred in nest boxes, so the reproductive success could easily be determined. Pellets, and feathers of bird prey species, were collected during the breeding seasons, and the food spectrum was determined based on these remains. There was no significant difference in the reproductive success of the kestrels between the three zones. Data on the number of fledged young indicated a sufficient food supply. In total, 9 species of mice and shrews, 23 bird species and 31 beetle species were identified as prey of kestrels. Urban kestrels specialise in hunting birds if mice and shrews are not readily available, with the house sparrow (Passer domesticus) as the favoured prey bird. Of note are anthropogenic food items, such as cutlet bones, that were found only in the city centre. This shows that the kestrel can adapt itself to humans with regard to its diet. There was no urban gradient with regard to reproductive success, but there was with the composition of food, such as the domination of bird prey in the city centre. The number of individual items of bird prey decreased from the centre to the outskirts. In conclusion, the results show that the kestrel is an opportunistic species which survives well anywhere in the city of Berlin.

Effect of felling on red fox (Vulpes vulpes) and pine marten (Martes martes) diets in transitional mixed forest in Belarus

In transitional mixed forests in northern and central Belarus the influence of intensified felling on the diets of red foxes Vulpes vulpes L. and pine martens Martes martes L. was investigated in two model forested terrains with sandy and clay top-grounds. A total of 1904 scats of red foxes and 1624 scats of pine martens were analysed over two periods differed by logging rate. When logging rate was conservative, red fox and pine marten diets were found to be similar, but under heavy logging feeding of the predator species shifted. In both model woodlands we found the same pronounced dietary trend of higher consumption of rodents, first of all, Microtus voles. The dietary changes were well related to the registered increase in Microtus vole numbers and total number of rodents in felling areas. The increased preying on rodents caused lower consumption of other food items, particularly medium-sized mammals (year-round) or/and birds or/and fruits (in the warm season) or/and mammalian carrion (in the cold season). In the conditions of intensified felling the food niches of the red fox and pine marten diverged mostly because of the great difference in the species structure of rodents consumed. Red foxes turned to preying on Microtus voles more frequently, but less on bank voles Myodes glareolus; while pine martens increased their taking of Microtus voles, continued foraging for bank voles and began taking slightly more of Apodemus mice. Before heavy logging dietary similarity between the red fox and pine marten was high and did not vary considerably through seasons and study areas, whereas after felling was intensified their diet overlap became lower.     

Diet breadth and overlap in three sympatric aerial insectivorous birds at the same location

Capsule We found significant differences in the composition and diversity of diet among House Martins, Barn Swallows and Swifts breeding in the same village in Poland.

Aims To evaluate the character and extent of diversity, specialization and overlap of diet between trophically similar nestlings of three species of aerial feeding birds breeding at the same location and differing considerably in foraging height.

Methods Diet was determined based on faecal analysis. Differences in composition and diversity of diet and food niche overlap were assessed through multivariate analysis of variance (manova), Shannon Diversity Index and the Pianka index. Diet specialization was measured by application of the Berger–Parker index of dominance.

Results manova indicated significant differences in diet composition among all three species. House Martins showed the most diverse diet, Swallows were intermediate and Swifts least diverse. Average body mass of all prey found in the diet of Swifts was nearly three times smaller than in Swallows and two times smaller than in House Martins.

Conclusion Our findings show that these three species consume the same types of insect prey, but they take different proportions, and hence biomass, of the major prey groups. House Martins had the widest niche and greatest overlap.     

Dietary shifts of sympatric buteos during a prey decline

Summary Diets of nesting Red-tailed Hawks (Buteo jamaicensis) and Ferruginous Hawks (Buteo regalis) were sampled before and after a decline in the hawks' principal prey species. Diets of pairs that shared their foraging ranges with interspecifics were contrasted with those of pairs whose home ranges did not overlap with interspecifics. Current theory predicts that diets should diverge during prey shortages and that overlap should be especially reduced in ranges shared with interspecifics. Species composition of the two hawks' diets diverged during the prey shortage, but the divergence was most pronounced in hawks that did not share foraging ranges with interspecifics. In contrast to predictions, the two species converged on similar-sized prey during the prey shortage. Available data on differences in habitat composition and prey availability at the sample sites did not explain the deviations from the predicted response. Although our findings on diet shifts differ from those of most other studies, the implications are consistent with observations of others who have studied assemblages of mammal-feeding raptors. We conclude that diet composition of the hawks we studied was not directly affected by interspecific competition and that shifts in diet overlap during prey shortages do not necessarily imply that interspecific competition is occurring.     

Habitat use by singing voles and tundra voles in the southern Yukon

Summary We investigated how far competitive interactions influence the use of habitats and relative abundance of two species of Microtus in the southwestern Yukon. We worked in the ecotone between alpine tundra and subalpine shrub tundra where populations of singing voles (Microtus miurus) and tundra voles (M. oeconomus) overlap little.We removed tundra voles from shrub tundra on one live-trapping area to look at the effect on the contiguous population of singing voles in alpine tundra. The removal of tundra voles did not affect the distribution or relative abundance of singing voles. The spatial distribution of these species and their movements within habitats suggest that they have a strong habitat preference.Populations of small mammals in the area are extremely dynamic and the relative importance of competitive interactions may change as density varies. At present we have no evidence that competition affects habitat use in M. miurus.     

Snow tracking reveals different foraging patterns of red foxes and pine martens

Red fox (Vulpes vulpes) shares similar prey preferences and co-occurs with several other carnivores, and is together with pine marten (Martes martes), the most common mesocarnivore in the northern boreal forest. Voles are important prey for both species, but it is unclear to what extent they compete for the same food resources in winter. Here, we use 2139 km and 533 km of meticulous snow tracking of red foxes and pine martens to evaluate their food niches. We measured hunting and digging behaviour, whether successful or not, and the effect of snow depth and temperature. Pine martens were restricted to forested habitats, whereas red foxes used a wide range of habitats. Red foxes were found to dig more often than pine martens, 0.67 vs. 0.39 digging events per kilometre. Hunting was less common and similar in both species, about 0.1 hunting event per kilometre. Pine martens were more efficient in hunting and finding food remains compared to red foxes. Increasing snow depth reduced hunting success and also reduced dig success of red foxes. Food niche overlap was small. Red foxes used mostly voles and carrion remains of ungulates, whereas pine martens used cached eggs and small birds. We suggest that caching eggs is an important strategy for pine martens to survive winter in northern latitudes. Snow depth was important for capturing voles, and thick snow cover appeared to mask the effect of vole peaks. Intensified land use, as clear-cutting and leaving slaughter remains from harvest, will benefit red foxes on the expense of pine martens. The ongoing climate change with warmer winters and less snow will likely further benefit the red fox.     

Investigating the influence of local weather conditions and alternative prey composition on the breeding performance of urban Eurasian Kestrels Falco tinnunculus

Capsule: Local weather conditions, but not a city-associated diet composition, influence the breeding performance of urban Eurasian Kestrels Falco tinnunculus.

Aims: We aim to explore the impact of diet composition, specifically a high proportion of avian prey, on the breeding performance of urban Eurasian Kestrels under different weather conditions.

Methods: The Eurasian Kestrel is known to exploit cities and occurs in the study area of Vienna, Austria (415?km²) at high breeding densities of 89–122 pairs/100?km². Unlike their rural conspecifics which specialize in voles, urban Kestrels are generalists, preying on passerines and mammals, but also reptiles and insects. We explore this alternative diet through pellet analyses along an urbanization gradient over a 5-year period and link diet to reproductive performance, taking local weather conditions into account.

Results: Our results showed that weather had a greater influence on breeding performance than did diet. Warm and dry weather during the arrival and courtship period was correlated with earlier egg-laying, higher nestling survival and an overall higher breeding success. Dry winters increased the proportion of mammalian prey, whereas low temperatures and high rainfall during the nestling phase increased the proportion of avian prey. Overall, a more diverse diet was also associated with higher rainfall, but only during the arrival and courtship period.

Conclusion: Our research shows a stronger connection between breeding performance and weather conditions than between breeding performance and diet composition, although there may be indirect effects of weather on prey availability which augment the impact of diet on reproductive output in the Eurasian Kestrel.     

Characterization of feeding habits,prey diversity and diet overlap of two sympatric species: Bronze catfish,Netuma bilineata (Valenciennes, 1840) and blacktip sea catfish,Plicofollis dussumieri (Valenciennes, 1840) in the northern Arabian Sea

This study describes the diet composition, diet overlap and prey diversity in two economically and ecologically important marine catfish species: the bronze catfish (Netuma bilineata) and the blacktip sea catfish (Plicofollis dussumieri) collected between January and December 2015 from the northern Arabian Sea coast of Pakistan. Stomach data of 342 bronze catfish and 283 blacktip sea catfish were examined. Diet composition and feeding strategy of each species by identification of stomach contents were weighed and enumerated. Both species primarily consumed teleosts, Brachyura and small amounts of a wide variety of prey items; however, specialization was a more common trait for the bronze catfish than for the black tip sea catfish. The Morisita‐Horn index showed considerable diet overlap in percentage of mean weights (0.847) than in numbers (0.612) between co‐prey pairs of both species. The highest Bray‐Curtis similarity between the southwest monsoon (SWM) and south inter monsoon (SIM) was observed in bronze catfish and lowest between autumn inter monsoon (AIM) seasons in both species. While Bray‐Curtis dissimilarity was observed, both species share diet niches and diet overlap. The present study evaluation of the feeding strategy and potential competition between two sympatric species of marine catfishes can provide the framework for their conservation and management in the region.     

Differences in time and space use between two sympatric Acrocephalus warblers with similar diets

Capsule: We found high diet overlap and different uses of space and time between Moustached Warblers Acrocephalus melanopogon and Reed Warblers Acrocephalus scirpaceus breeding in sympatry at a marshland in Spain.

Aims: To study the degree of diet overlap between both species, their space use on a local scale and their breeding phenologies.

Methods: We studied the breeding phenologies of the two species by standardized ringing activity. Spatial distribution was investigated by point counts. We determined diet composition from emetic samples and we collected invertebrates by standardized sweep-netting to estimate food availability.

Results: Diet and prey selection were similar among species. Conversely, spatial overlap was relatively small (<50%) and breeding phenologies were not synchronized. Both food availability and the overall abundance of the two species increased throughout the breeding season.

Conclusion: The two species are potential competitors for food and the observed differences in spatial and temporal niches may represent a way to lower competition for trophic resources: Moustached Warblers could reduce competition by breeding early, while Reed Warblers could avoid settling in areas occupied by the other species.     

Marine copepod diversity patterns and the metabolic theory of ecology

The ongoing climate change has improved our understanding of how climate affects the reproduction of animals. However, the interaction between food availability and climate on breeding has rarely been examined. While it has been shown that breeding of boreal birds of prey is first and foremost determined by prey abundance, little information exists on how climatic conditions influence this relationship. We studied the joint effects of main prey abundance and ambient weather on timing of breeding and reproductive success of two smaller (pygmy owl Glaucidium passerinum and Tengmalm’s owl Aegolius funereus) and two larger (tawny owl Strix aluco and Ural owl Strix uralensis) avian predator species using long-term nation-wide datasets during 1973–2004. We found no temporal trend either in vole abundance or in hatching date and brood size of any studied owl species. In the larger species, increasing late winter or early spring temperature advanced breeding at least as much as did high autumn abundance of prey (voles). Furthermore, increasing snow depth delayed breeding of the largest species (Ural owl), presumably by reducing the availability of voles. Brood size was strongly determined by spring vole abundance in all four owl species. These results show that climate directly affects the breeding performance of vole-eating boreal avian predators much more than previously thought. According to earlier studies, small-sized species should advance their breeding more than larger species in response to increasing temperature. However, we found an opposite pattern, with larger species being more sensitive to temperature. We argue that this pattern is caused by a difference in the breeding tactics of larger mostly capital breeding and smaller mostly income breeding owl species.     

Diet composition of two larval headwater stream salamanders and spatial distribution of prey

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Diet composition and food consumption of the grey heron (Ardea cinerea) from breeding colonies in northern Poland

The diet of the grey heron (Ardea cinerea) was investigated in three colonies in northern Poland (Mosty, Kiersity and Kty Rybackie) varying in size, habitat type and composition of feeding areas used by birds. The diet was assessed during the breeding season on the base of pellets and regurgitated food. Pellets from all studied colonies consisted mainly of mammal hair (almost 100% of samples), bone remains (20–24%) and invertebrate remains (26–51%, mainly water beetle, Dytiscus marginalis). Bones and hair of water vole (Arvicola terrestris) and Microtus voles were the most common remains of mammals recorded in pellets. Remains of other animal taxa were found sporadically. In the colonies at Mosty and Kiersity, fish composed more than 95% of collected regurgitated prey items. Food consumption of herons from the biggest colony at Kty Rybackie was estimated to exceed 100 t per season. Herons from Mosty consumed ca 46–52 t, from Kiersity 30–38 t of food per season. The impact of herons on fish communities was much lower than other losses due to predation (herons from Kty Rybackie caught only 6% of fishermen bycatch, birds from Mosty took 10–14% of round gobies eaten by cormorants).