Numerical and functional responses of Common Buzzards Buteo buteo to prey abundance on a Scottish grouse moor

Predators will often respond to reductions in preferred prey by switching to alternative prey resources. However, this may not apply to all alternative prey groups in patchy landscapes. We investigated the demographic and aggregative numerical and functional responses of Common Buzzards Buteo buteo in relation to variations in prey abundance on a moor managed for Red Grouse Lagopus lagopus scotica in south‐west Scotland over three consecutive breeding and non‐breeding seasons. We predicted that predation of Red Grouse by Buzzards would increase when abundance of their preferred Field Vole Microtus agrestis prey declined. As vole abundance fluctuated, Buzzards responded functionally by eating voles in relation to their abundance, but they did not respond demographically in terms of either breeding success or density. During a vole crash year, Buzzards selected a wider range of prey typical of enclosed farmland habitats found on the moorland edge but fewer Grouse from the heather moorland. During a vole peak year, prey remains suggested a linear relationship between Grouse density and the number of Grouse eaten (a Type 1 functional response), which was not evident in either intermediate or vole crash years. Buzzard foraging intensity varied between years as vole abundance fluctuated, and foraging intensity declined with increasing heather cover. Our findings did not support the prediction that predation of Red Grouse would increase when vole abundance was low. Instead, they suggest that Buzzards predated Grouse incidentally while hunting for voles, which may increase when vole abundances are high through promoting foraging in heather moorland habitats where Grouse are more numerous. Our results suggest that declines in their main prey may not result in increased predation of all alternative prey groups when predators inhabit patchy landscapes. We suggest that when investigating predator diet and impacts on prey, knowledge of all resources and habitats that are available to predators is important.     

Field Vole Microtus agrestis abundance and Hen Harrier Circus cyaneus diet and breeding in Scotland

In many parts of the global range, voles form an important part of the diet of Hen Harriers Circus cyaneus , and breeding numbers are correlated with the abundance of these small mammals. In Scotland, however, little information is available on harrier diet in the spring and our understanding of causes of variation in harrier breeding density is complicated by human interference. In this paper we explore the relationship between Field Vole Microtus agrestis abundance and harrier spring diet, density and productivity in southern Scotland. Over three years, voles occurred on average in 67% of pellets, and 79% in years of high and intermediate vole abundance. From 1992, the number of breeding harriers increased following protection from illegal persecution. After accounting for this trend, harrier numbers correlated strongly with vole abundance. Harrier clutch size was also correlated with vole abundance. Although fledging success tended to be greater in years of vole increase than in years of vole decline, fledging success was not significantly correlated with the relative abundance of voles, or with the abundance of Meadow Pipits or Red Grouse chicks.     

The effect of oilseed rape occurrence on main prey abundance and breeding success of the Common Buzzard Buteo buteo

Capsule The occurrence of oilseed rape increased main prey abundance and breeding success of Common Buzzards.

Aims We tested whether the occurrence of oilseed rape influences the abundance of Common Voles, i.e. the main prey of Common Buzzards and so also nesting activity and breeding success of Common Buzzards.

Methods The study was carried out in 2005–2012 in a 38?km² area in western Poland, where oilseed rape plantations (12–106?ha) covered 18% of the agricultural land. The number of active burrow entrances was used as an index of vole abundance in various crops, and Buzzard breeding performance, i.e. the occurrence of annual nesting attempts in individual long-term nesting sites as well as the presence and number of fledglings, was estimated by observations of their nests.

Results The index of vole abundance was highest in oilseed rape, and judging by the proportion of active burrow entrances (33–77%), the plantations of rape typically supported a larger portion of the local vole population than other crops. The acreage of oilseed rape fields around individual nesting sites of Buzzards did not affect the probability of nesting attempts in these sites. However, the probability of successful nesting and the number of fledglings per successful nest increased with the area of oilseed rape around the Buzzard nesting sites.

Conclusion The occurrence of oilseed rape may positively affect prey availability and in turn the breeding success of Buzzards. The spread of oilseed rape may therefore also be beneficial for other vole-eating raptors hunting in the agricultural landscapes.     

Dietary shifts,niche relationships and reproductive output of coexisting Kestrels and Long-eared Owls

Summary Food samples of breeding Kestrels (Falco tinnunculus) and Long-eared Owls (Asio otus) were collected in the peak and low phase of their preferred prey (Microtus voles) in western Finland. Diets of pairs that bred as neighbours (1 km) with interspecifics were compared with those of non-neighbours. In both species, neighbouring pairs fed less on Microtus voles and more on alternative prey than did non-neighbours. Competition theory predicts that diet overlap should be lower during prey shortage and that diet similarity should be especially reduced in neighbouring pairs. Observations were consistent with expectations: diet similarity was lower in the low vole years and neighbouring pairs showed less diet overlap that non-neighbours. Differences in habitat composition and prey availability at the sample sites should not confuse the results. In addition to the high diet similarity, hunting habitats and nest sites of the species overlapped almost completely; they only showed clear temporal segregation in hunting. Probably because of food competition, the neighbouring pairs of both species produced significantly fewer young than the non-neighbours. These results contrast with the view that the diet composition and dietary shift of rodent-feeding predatory birds can be interpreted in terms of simple opportunistic foraging. In the breeding season, interspecific competition for food seems to be an important factor that affects the niches of these species, especially in northern areas, where the seasonal low phase of voles in spring and the number of alternative prey are lower than in more southern areas.     

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.     

Diet specialisation and foraging efficiency under fluctuating vole abundance: a comparison between generalist and specialist avian predators

Specialist species, using a narrow range of resources, are predicted to be more efficient when foraging on their preferred food than generalist species consuming a wider range of foods. We tested whether the foraging efficiency of the pallid harrier Circus macrourus, a vole specialist, and of sympatric Montagu's harriers C. pygargus, a closely related generalist, differed in relation to inter‐annual variations in vole abundance over five years (including two peak‐ one intermediate and two low vole abundance years). We show that the hunting parameters of pallid harriers strongly varied with vole abundance (higher encounter rates, capture rates and proportion of successful strikes in high than intermediate and low vole abundance years, respectively), whereas Montagu's harriers showed stable capture rates and hunting success (proportion of strikes that were successful), irrespective of vole abundance. Encounter rates and capture rates were higher for pallid than for Montagu's harriers when voles were abundant, but lower when voles were scarce. The hunting success of pallid harriers was also lower than that of Montagu's harriers when voles were scarce, and when they had to target alternative preys, in particular birds. Overall, estimated biomass intake rate was 40% higher for pallid harriers than for Montagu's harriers when voles were abundant, but 50% lower when voles were scarce. Our results indicate that specialists predators, like pallid harriers, which evolve specific adaptations or breeding strategies, do better when their preferred prey is abundant, but may face a cost of specialisation, being not efficient enough when their preferred prey is scarce. These results have broader implications for understanding why specialist predators are, in general, more vulnerable than generalists, and for predicting how specialists can cope with rapid environmental changes affecting the abundance or predictability of their preferred resources.     

Prey abundance and reproductive success of the golden eagle Aquila chrysaetos in Sweden

Reproductive output of the golden eagle Aquila chrysaetos (L.), was studied in two areas within the species distribution in northern Sweden during 1975–1980. Reproductive success was compared with the abundance (based on hunting bag statistics) of small game prey species and with the microtine cycles. The proportion of golden eagle pairs with successful breeding and the number of young produced per occupied territory varied greatly between years (21–85% and 0.27–1.24, respectively). In the northern study area I found a significant correlation between the proportion of pairs with successful breeding versus total hunting bag of small game species. There were also significant correlations between vole density and breeding success one year later. This was not the case in the southern study area, mainly due to a good reproductive year for golden eagles in 1977 when small game species were scarce. The good reproductive output in 1977 may be explained by favourable weather conditions that spring. Whether breeding occurs or not is probably determined by prey abundance early in spring just before the golden eagle female lays her eggs.     

Long-term population declines in Danish trans-Saharan migrant birds

Capsule Long-distance migrant birds show less favourable trends than sedentary/short-distance species.

Aims To use breeding bird surveys to contrast population trends amongst common species according to their migration pattern.

Methods Changes in abundance of 62 Danish breeding sedentary, short-distance (Europe/North Africa) or long-distance (trans-Saharan) migrants were described by fitting log linear regression models to point-count census data gathered during 1976–2005.

Results Trans-Saharan migrants declined by 1.3% per annum during this period, while short-distance migrants and sedentary species increased by 1.4% and 1.0% per annum, respectively. There were no significant decadal declines amongst species using different summer breeding habitats, except for wetlands, and there was no consistent variation in trends associated with wintering regions or habitats or diet.

Conclusions More information is urgently needed on diet, feeding ecology, habitat requirements, winter distribution and intra-African movements of the commoner European summer visitors to identify causes of the declines and highlight when in the annual cycle detrimental effects occur. Studies linking individuals on their breeding, staging and wintering grounds are especially needed. Danish trends resemble those from elsewhere in Europe, confirming that restoration to favourable conservation status requires inter-continental action to meet European and global targets to reduce or halt biodiversity loss.     

The difference between generalist and specialist: the effects of wide fluctuations in main food abundance on numbers and reproduction of two co-existing predators

Specialist individuals within animal populations have shown to be more efficient foragers and/or to have higher reproductive success than generalist individuals, but interspecific reproductive consequences of the degree of diet specialisation in vertebrate predators have remained unstudied. Eurasian pygmy owls (hereafter POs) have less vole-specialised diets than Tengmalm's owls (TOs), both of which mainly subsist on temporally fluctuating food resources (voles). To test whether the specialist TO is more limited by the main prey abundance than the generalist PO, we studied breeding densities and reproductive traits of co-existing POs and TOs in central-western Finland during 2002–2019. Breeding densities of POs increased with augmenting densities of voles in the previous autumn, whereas breeding densities of TOs increased with higher vole densities in both the previous autumn and the current spring. In years of vole scarcity, PO females started egg-laying earlier than TOs, whereas in years of vole abundance TO females laid eggs substantially earlier than PO females. The yearly mean clutch size and number of fledglings produced of both POs and TOs increased with abundance of voles in the current spring. POs laid large clutches and produced large broods in years of both high and low vole abundance, whereas TOs were able to do so only in years of high vole abundance. POs were able to raise on average 73% of the eggs to fledglings whereas TOs only 44%. The generalist foraging strategy of POs including flexible switching from main prey to alternative prey (small birds) appeared to be more productive than the strictly vole-specialized foraging strategy of TOs. In contrast to earlier studies at the individual-level, specialist predators at the species level (in this case TOs) appear to be less effective than generalists (POs), but diet specialisation was particularly costly under conditions when scarcity of main foods limited offspring production.     

Is female investment in defence correlated with breeding success in Hen Harriers Circus cyaneus

CapsuleThe intensity of parental defence, irrespective of the value of offspring, may be one of the proximate causes of breeding success, reflecting the quality of breeders.

Aims To test whether female Hen Harrier investment in defence towards a human predator influences breeding success.

Methods Generalized linear mixed models were used, with nest content, date of visit, the interaction (nest content*date), breeding success (number of fledglings/clutch size) and presence or absence of male as explanatory variables. ‘Female’ was included as a random variable in the model. Alarm call rate by a female Hen Harrier during the first three minutes after my arrival at a nest was used as a dependent variable (as an indicator of parental investment in defence towards a human predator).

Results Females were present and alarmed in 100% of visits whereas males were present in 37% of visits. The individual variations in female investment in defence, after controlling for the principal determinants of nest defence, were significantly correlated with breeding success of each pair.

Conclusion The relationship between female investment in nest defence and reproductive success can be explained through differences in female quality. Good quality females, as measured by the level of investment in nest defence, probably also invested more in other breeding aspects not controlled in this study obtaining higher reproductive success.     

Pesticides and birds—A review of the situation in Great Britain in 1965

Capsule Arrival date strongly influenced date of breeding and breeding success.

Aim To check our hypothesis that in years with low April temperatures, i.e. when storks started to breed, the relationship between timing of breeding and success would be absent, meaning that migrants would have no advantage by returning early to their breeding area.

Methods We collected data in Poznań province, western Poland during the period 1983–2002. Based on local climatological data we selected eight cold spring years and compared them with the 12 ‘normal’ years. We analysed how weather affected the timing and arrival pattern of White Stork.

Results The two groups of years did not differ significantly in population size, but in normal years the arrival date of both parents was earlier. Arrival date strongly influenced (was positively correlated with) date of breeding and (negatively correlated with) breeding success.

Conclusion The slopes describing the above relationships did not differ significantly between the two groups of years. Therefore, we speculate that natural selection strongly favours birds that return early from the wintering grounds.     

Contrasting phenotypic correlations in food provision of male Tengmalm's owls (Aegolius funereus) in a temporally heterogenous environment

Summary We examined the food provision rate of male Tengmalm's owls,Aegolius funereus, during one 3 year vole cycle consisting of consecutive low, increase and peake vole years. The data were collected in the midnestling period when males provisioned the whole family. In the low vole year, males with a low loading index (g/cm²) of flying area fed their offspring more often than did males with a high loading index, whereas in the peak vole year the opposite trend was evident. Similar relationships were found in the food mass provisioned to the nest. In the increase vole year, male body size had no effect on feeding efficiency. In the peak vole year, when large voles are abundant, heavy males preyed on larger voles than were generally available in their territories, indicating that largeness may increase strike power in hunting attempts. In the low vole year, when breeding is costly due to food scarcity and extensive hunting area, small males are more economical fliers and efficient hunters than large males. The contrasting trends in correlations between male size and feeding efficiency in years of vole abundance versus scarcity suggest that no fixed phenotype may most efficiently cope with variable food supply.     

Effects of individual life-history traits and weather on reproductive output of Black-headed Gulls Larus ridibundus breeding in the Wadden Sea, 1991–97

Capsule Annual breeding success was relatively constant and is shown to be related to clutch size and growth rate and to be positively affected by rain during egg-laying and advanced chick-rearing phase.

Aims To provide the first long-term data on breeding success of Black-headed Gulls in the Wadden Sea, to analyse its intrinsic and environmental co-variates, and to re-assess the significance of the severe winter 1995/96 on reproduction.

MethodsIn a relatively small colony at the Wadden Sea coast, clutches were selected randomly and enclosed to determine clutch size, egg biometrics, hatching and breeding success, and chick development in 1991 and from 1994 to 1997. Weather data (temperature, rain, wind) were related to life-history traits (clutch size, egg volume, chick development) and reproductive success.

Results Mean annual breeding success was 0.7 fledglings per pair. In 1996, breeding success tended to be lower and chick growth rate was significantly lower. Hatching success was lower in small clutches, chick mortality increased with decreasing chick growth rate. Rain during the egg-laying phase increased clutch size and decreased clutch predation. Rain during the early post-natal phase impaired chick growth. Rain during the phase of linear growth affected chick growth and fledging success positively and brood predation negatively.

Conclusions Breeding success of Black-headed Gulls breeding in the Wadden Sea is relatively constant between years, probably due to the use of terrestrial and marine feeding habitats. Rain may increase the availability of intertidal and terrestrial prey and thus may affect time budgets and food provisioning of parents positively. Lower breeding success in 1996 might have been caused by a relatively dry breeding season and possibly by the preceding severe winter.     

Numerical responses of an avian predator to prey fluctuations in a temperate latitude: breeders vs. entire population

Distinct numerical responses of predators to fluctuations in the abundance of their prey are often observed in northern regions but occur more rarely in temperate latitudes. This statement is, however, mostly based on observations of breeding populations, while in some predators, for example in raptors, numerous non-breeding floaters can occur. I estimated the breeding density and reproductive performance (nest survey) as well as the density of entire population (transects with distance sampling) of the common buzzard Buteo buteo in western Poland (52°N) in the years 2005–2014 to test the hypotheses that in temperate latitudes: (1) the breeding population of these birds does not show any numerical response, understood as annual changes in their abundance; (2) its reproductive success, however, changes with the abundance of main prey, the common vole Microtus arvalis; and (3) hence the entire buzzard population (including potential immature floaters) present in a given area during the nesting period responds numerically with some time delay. The reproductive success of buzzards was positively correlated with their prey abundance. Contrary to my predictions, however, the breeding population of buzzards showed a slight numerical response with a 3-year lag and the entire population tracked vole fluctuations without any time delay. The immediate numerical response of the entire buzzard population was probably caused by large-scale movements of floaters. Such rapid numerical responses of some predators may contribute to the relative stability of prey populations in temperate latitudes compared to northern regions.     

Effect of trophic status of a deep-water lake on breeding Great Crested Grebes Podiceps cristatus during a phase of recovery from eutrophication: a long-term study

Capsule: Trophic status of a deep-water lake was the main driver of changes in breeding population size of Great Crested Grebes Podiceps cristatus while reproductive success was also strongly affected by weather parameters.

Aims: To determine the effects of changes in nutrient status of a formerly highly-eutrophicated deep-water lake and other environmental parameters on a Great Crested Grebe population during a phase of re-oligotrophication.

Methods: Annual surveys were carried out on a natural lake in Switzerland over a period of 25 years to determine breeding population size and reproductive success. The effects of phosphorus content, other limnological parameters and weather variables were analysed with quasi-Poisson models.

Results: The breeding population increased from 80 pairs in 1992 to 417 pairs in 2001, after which numbers showed strong fluctuations. Total phosphorus content in the lake had a strong negative effect on breeding population size. A significant positive correlation was found with the national population index. Reproductive success fluctuated strongly but showed an overall decline. The model indicated positive effects on reproductive success of phosphorus and negative effects of the number of days with strong wind. Rapid water-level increases in early summer and water transparency in June led to higher proportions of late broods.

Conclusion: Phosphorus concentration was identified as the main driver affecting the breeding population of Great Crested Grebes during the phase of recovery of the lake from a highly-eutrophic state. Results indicate that mesotrophic to eutrophic conditions enabled a large population and high breeding success. Reproductive output was further negatively affected by strong wind during a critical breeding phase.     

Parental provisioning behaviour in Pied Flycatchers Ficedula hypoleuca is well adjusted to local conditions in a mosaic of deciduous and coniferous habitat

Capsule Pied Flycatchers are better able than Great Tits to adjust their feeding behaviour to varying conditions in the same area.

Aims Great Tits breeding in a mosaic of deciduous and coniferous forests in the northern temperate region exhibit consistently lower breeding success in their preferred deciduous habitat than in coniferous habitat. This was explained by the unexpectedly poor nestling feeding conditions in deciduous forests of this region. We studied whether the same paradox applies to Pied Flycatchers that occupy the same habitats in the same area.

Methods Parental provisioning behaviour was studied using video‐recording and experimental manipulation. Caterpillar abundance and basic breeding parameters were measured in different habitat types.

Results Parental provisioning frequency and the proportion of caterpillars in nestling diet was lower, while food objects were on average larger, in coniferous compared with deciduous habitat. However, the total volume of caterpillars and adult Lepidoptera delivered to nestlings did not differ between habitats. In contrast to Great Tits, offspring body parameters in Pied Flycatchers did not differ between habitat types.

Conclusions These results demonstrate how the relative suitability of particular habitat types varies between species and is dependent upon geographical location.     

Predator control and the density and reproductive success of grouse populations in Finland

The impact of predator removal or protection on the reproductive success and density of grouse populations was studied experimentally in southern and northern Finland. In predator removal areas small and medium-sized carnivores wore efficiently hunted, while in predator protection areas hunting was prohibited. Both predator (red fox, raccoon dog, pine marten and stoat) and grouse populations were monitored. The breeding success of grouse, indicated by the young/adults ratio, decreased in the predator protection area in northern Finland during a vole low. Despite a decline in the vole population, the breeding success of grouse did not decline in the predator removal area. The mean brood size during the experiment was significantly higher in the removal areas than in the protection areas both in southern and northern Finland. Predator removal/protection thus affected the reproductive success of grouse, but the impact of control on adult grouse populations was not as evident.     

Week-end bias at observatories

Capsule Although subject to human disturbance Turtle Doves do nest successfully in these olive and orange orchards.

Aim To investigate the breeding ecology of Turtle Doves in a man‐made agricultural habitat in central Morocco.

Methods Turtle Dove nests were monitored in orange and olive orchards over three years (2006, 2007 and 2008). Nest abundance, nest location, egg‐laying chronology, clutch size, nest survival rates and breeding success were determined and compared between orchard types.

Results The Moroccan population of Turtle Doves start breeding earlier than European populations. Clutch size, nest survival rates and breeding success were similar in orange and olive orchards. Nest location differed between orange and olive trees. Nest densities were 16 nests/ha in olive orchards and 45 nests/ha in orange orchards. Nest success rate averaged 48%. Daily nest survival rates did not vary according to orchard types, year and date. In the two orchards, no nest position variables were significant predictors of nesting success.

Conclusion Although highly frequented by people, fruit orchards seem to be suitable breeding habitats for Turtle Doves in this region.     

Effects of vole fluctuations on the population dynamics of the barn owl <Emphasis Type="Italic">Tyto alba</Emphasis>

Many predator species feed on prey that fluctuates in abundance from year to year. Birds of prey can face large fluctuations in food abundance i.e. small mammals, especially voles. These annual changes in prey abundance strongly affect the reproductive success and mortality of the individual predators and thus can be expected to influence their population dynamics and persistence. The barn owl, for example, shows large fluctuations in breeding success that correlate with the dynamics in voles, their main prey species. Analysis of the impact of fluctuations in vole abundance (their amplitude, peaks and lows, cycle length and regularity) with a simple predator prey model parameterized with literature data indicates population persistence is especially affected by years with low vole abundance. In these years the population can decline to low owl numbers such that the ensuing peak vole years cannot be exploited. This result is independent of the length and regularity of vole fluctuations. The relevance of this result for conservation of the barn owl and other birds of prey that show a numerical response to fluctuating prey species is discussed.     

The effect of extreme weather events on breeding parameters of the White Stork Ciconia ciconia

Capsule Although the White Stork avoids adverse weather conditions by modifying its arrival and breeding, it cannot avoid extreme weather events during the breeding season.

Aims To show how extreme weather conditions can influence breeding attempts of a large, long-lived species, the White Stork.

Methods We analysed data on arrivals of White Storks in Western Poland from 2005 to 2013 and detailed breeding biology parameters from 2009 to 2013 in relation to weather conditions. We analysed breeding success and breeding failure rate from 1974 to 2013.

Results In years with a cold March White Storks arrived later than when March was warmer. Frost during incubation negatively influenced the hatching success. Extreme weather events caused high late mortality even for nestlings older than 30 days. Data from 27 breeding seasons showed a significant increase in mean breeding success but also a significant increase in the proportion of pairs which lost broods in the nestling stage.

Conclusion The White Stork can modify its arrival in response to current weather conditions on the breeding grounds but it cannot respond to extreme weather events. Due to increasing frequency of extreme weather events caused by climate change, White Stork breeding success may decrease in the future.