Long-term consistency in susceptibility of prey species to predation by an avian predator.

Selection by predators affects prey through competition for limiting resources. This not only has consequences for direct mortality but also indirectly affects disturbance. Changes in the intensity of selection on prey by predators may affect the size of prey populations, with consequences for their short- or long-term interactions. We assessed whether predation by northern goshawks Accipiter gentilis modified the composition of prey communities consistently along a temporal gradient, showing long-term consistency in susceptibility of prey species to predation. We followed six populations of the goshawk in two biomes in Denmark and Finland during 1949-2019. Susceptibility to goshawk predation in 2005-2017 in Denmark was only weakly related to susceptibility to goshawk predation in 1977-2004. In Finland, susceptibility of shared prey species to goshawk predation was positively related between periods. The average difference in susceptibility to goshawk predation between periods was considerably higher in Denmark than in Finland. Susceptibility of prey species to predation in goshawks increased with latitude and body mass of prey species, and decreased with period of time and population density of prey species. The changes in susceptibility to predation suggest changes in the characteristics of the local prey pools.

The abundance of small mammals is positively linked to survival from nest depredation but negatively linked to local recruitment of a ground nesting precocial bird.

Generalist predators using small mammals as their primary prey are suggested to shift hunting alternative prey such as bird nests, when small mammals are in short supply (the alternative prey hypothesis, APH). Nest survival and survival of young individuals should be positively linked to small mammal abundance and negatively linked to predator abundance, but little information exists from survival of chicks, especially until recruitment. We test these predictions of the APH using 13 years (2002-2014) of life history data from a ground nesting shorebird breeding on coastal meadows. We use small mammal abundance in the previous autumn as a proxy for spring predator abundance, mainly of mammalian predators. We examine whether small mammal abundance in the spring and previous autumn explain annual variation in nest survival from depredation and local recruitment of the southern dunlin Calidris alpina schinzii. As predicted by the APH, survival from nest predation was positively linked to spring small mammal abundance and negatively linked to autumn small mammal abundance. Importantly, local recruitment showed opposite responses. This counterintuitive result may be explained by density-dependent survival. When nest depredation rates are low, predators may show stronger numerical and functional responses to high shorebird chick abundance on coastal meadows, whereas in years of high nest depredation, few hatching chicks lure fewer predators. The opposite effects on nest and local recruitment demonstrate the diverse mechanisms by which population size variation in primary prey can affect dynamics of alternative prey populations.

Breeding near heterospecifics as a defence against brood parasites: can redstarts lower probability of cuckoo parasitism using neighbours?

Breeding habitat choice based on the attraction to other species can provide valuable social information and protection benefits. In birds, species with overlapping resources can be a cue of good quality habitats; species with shared predators and/or brood parasites can increase joint vigilance or cooperative mobbing, while raptors may provide a protective umbrella against these threats. We tested whether the migratory common redstart (Phoenicurus phoenicurus) is attracted to breed near active nests of the great tit (Parus major), a keystone-information source for migrant passerine birds, or a top predator, the northern goshawk (Accipiter gentilis). This system is unique to test these questions because the redstart is a regular host for the common cuckoo (Cuculus canorus). Therefore, we also evaluated other possible benefits coming from the heterospecific attraction, especially in terms of reducing brood parasitism risk. We monitored redstart occupancy rates, onset of breeding, reproductive investment, and followed nest outcomes in terms of brood parasitism, nest predation risk and overall reproductive success. Redstarts avoided breeding near goshawks, but showed neither attraction nor avoidance to breed next to great tits. Both neighbours neither reduced brood parasitism risk nor affected overall nesting success in redstarts. Redstarts may not use heterospecific attraction for settlement decisions, as associations with other species can only exist when some benefits are gained. Thus, environmental cues may be more important than social information for redstarts when breeding habitat choice. Other front-line defence strategies may have a better impact reducing breeding negative interactions, such brood parasitism.

Comparison of Nest Defense Behaviors of Goshawks (Accipiter gentilis) from Finland and Montana.

As human impacts on wildlife have become a topic of increasing interest, studies have focused on issues such as overexploitation and habitat loss. However, little research has examined potential anthropogenic impacts on animal behavior. Understanding the degree to which human interaction may alter natural animal behavior has become increasingly important in developing effective conservation strategies. We examined two populations of northern goshawks (Accipiter gentilis) in Montana and Finland. Goshawks in Finland were not protected until the late 1980s, and prior to this protection were routinely shot, as it was believed that shooting goshawks would keep grouse populations high. In the United States, Goshawk were not managed as predator control. Though aggressive nest defense has been characterized throughout North America, goshawks in Finland do not show this same behavior. To quantify aggression, we presented nesting goshawks with an owl decoy, a human mannequin, and a live human and recorded their responses to each of the trial conditions. We evaluated the recordings for time of response, duration of response, whether or not an active stimulus was present to elicit the response (i.e., movement or sound), and the sex of the bird making the response. We used t-Test with unequal variance to compare mean number of responses and response duration. Our results suggested that goshawks in Montana exhibit more aggressive nest defense behaviors than those in Finland. While this could be due to some biotic or abiotic factor that we were not able to control for in a study on such a small scale, it is also possible that the results from this study suggest another underlying cause, such as an artificial selection pressure created by shooting goshawks.

Degradation in landscape matrix has diverse impacts on diversity in protected areas.

INTRODUCTION: A main goal of protected areas is to maintain species diversity and the integrity of biological assemblages. Intensifying land use in the matrix surrounding protected areas creates a challenge for biodiversity conservation. Earlier studies have mainly focused on taxonomic diversity within protected areas. However, functional and especially phylogenetic diversities are less studied phenomena, especially with respect to the impacts of the matrix that surrounds protected areas. Phylogenetic diversity refers to the range of evolutionary lineages, the maintenance of which ensures that future evolutionary potential is safeguarded. Functional diversity refers to the range of ecological roles that members of a community perform. For ecosystem functioning and long-term resilience, they are at least as important as taxonomic diversity. AIM: We studied how the characteristics of protected areas and land use intensity in the surrounding matrix affect the diversity of bird communities in protected boreal forests. We used line-transect count and land-cover data from 91 forest reserves in Northern Finland, and land-cover data from buffer zones surrounding these reserves. We studied if habitat diversity and productivity inside protected areas, and intensity of forest management in the matrix have consistent effects on taxonomic, functional and phylogenetic diversities, and community specialization. RESULTS: We found that habitat diversity and productivity inside protected areas have strong effects on all diversity metrics, but matrix effects were inconsistent. The proportion of old forest in the matrix, reflecting low intensity forest management, had positive effects on community specialization. Interestingly, functional diversity increased with increasing logging intensity in the matrix. CONCLUSIONS: Our results indicate that boreal forest reserves are not able to maintain their species composition and abundances if embedded in a severely degraded matrix. Our study also highlights the importance of focusing on different aspects of biodiversity.

Diet shift induced rapid evolution of size and function in a predatory bird.

A predator's body size correlates with its prey size. Change in the diet may call for changes in the hunting mode and traits determining hunting success. We explored long-term trends in sternum size and shape in the northern goshawk by applying geometric morphometrics. Tetraonids, the primary prey of the goshawk, have decreased and been replaced by smaller birds in the diet. We expected that the size of the goshawk has decreased accordingly more in males than females based on earlier observations of outer morphology. We also expected changes in sternum shape as a function of changes in hunting mode. Size of both sexes has decreased during the preceding decades (1962-2008), seemingly reflecting a shift in prey size and hunting mode. Female goshawks hunting also mammalian prey tend to have a pronouncedly "Buteo-type" sternum compared to males preying upon birds. Interestingly, the shrinkage of body size resulted in an increasingly "Buteo-type" sternum in both sexes. In addition, the sternum shape in birds that died accidentally (i.e., fit individuals) was more Buteo-type than in starved ones, hinting that selection was towards a Buteo-type sternum shape. We conclude that these observed patterns are likely due to directional selection driven by changes in the diet towards smaller and more agile prey. On the other hand, global warming is predicted to also cause a decrease in size, thus these two scenarios are inseparable. Because of difficulties in studying fitness-related phenotypic changes of large raptors in the field, time series of museum exemplars collected over a wide geographical area may give answers to this conundrum.

Coupling in goshawk and grouse population dynamics in Finland.

Different prey species can vary in their significance to a particular predator. In the simplest case, the total available density or biomass of a guild of several prey species might be most relevant to the predator, but behavioural and ecological traits of different prey species can alter the picture. We studied the population dynamics of a predator-prey setting in Finland by fitting first-order log-linear vector autoregressive models to long-term count data from active breeding sites of the northern goshawk (Accipiter gentilis; 1986-2009), and to three of its main prey species (1983-2010): hazel grouse (Bonasa bonasia), black grouse (Tetrao tetrix) and capercaillie (T. urogallus), which belong to the same forest grouse guild and show synchronous fluctuations. Our focus was on modelling the relative significance of prey species and estimating the tightness of predator-prey coupling in order to explain the observed population dynamics, simultaneously accounting for effects of density dependence, winter severity and spatial correlation. We established nine competing candidate models, where different combinations of grouse species affect goshawk dynamics with lags of 1-3 years. Effects of goshawk on grouse were investigated using one model for each grouse species. The most parsimonious model for goshawk indicated separate density effects of hazel grouse and black grouse, and different effects with lags of 1 and 3 years. Capercaillie showed no effects on goshawk populations, while the effect of goshawk on grouse was clearly negative only in capercaillie. Winter severity had significant adverse effects on goshawk and hazel grouse populations. In combination, large-scale goshawk-grouse population dynamics are coupled, but there are no clear mutual effects for any of the individual guild members. In a broader context, our study suggests that pooling data on closely related, synchronously fluctuating prey species can result in the loss of relevant information, rather than increased model parsimony.

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

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

Competitive interactions among raptors in boreal forests.

We examined inter-specific interactions among goshawks ( Accipiter gentilis), common buzzards (Buteo buteo) and honey buzzards (Pernis apivorus) in western Finland in 1983-1996. Because goshawks are among the largest birds of prey species in boreal forests they may take over the nest of smaller and less-competitive forest-dwelling raptors when searching for suitable places for breeding. Accordingly, more than half of newly established goshawk territories were found on the territories previously occupied by the common buzzard and the honey buzzard. Otherwise, territory sharing between these species was rare. Fledgling production of honey buzzards was not associated with the presence of goshawks, probably owing to the almost 2 months later onset of breeding. This probably decreases competitive interactions between these two species. An intensive interference competition, instead, seemed to be evident between common buzzards and goshawks, because the fledgling production of common buzzards was decreased by 20% as a result of failures during incubation and nestling period in the vicinity (<1 km) of occupied goshawk nests. Similarly, territory occupancy of common buzzards till the next breeding season was significantly reduced in the presence of goshawks. Relatively high proportions of occupied buzzard territories (17%) in the study area were shared by breeding goshawks on the same territory. This suggests that although their diets are dissimilar they inhabit similar habitats and might compete for the available prime nesting habitats within forest landscapes. In addition, goshawks benefit from taking over the complete nests of other raptors, imposing upon the original owners of the nest, because building a large stick nest is probably energetically costly. As a large raptor, the goshawk apparently has a competitive advantage over smaller ones, and may have an ever-increasing impact on smaller birds of prey, if there is a lack of sheltered forests inducing competition for the available nest sites.