Interactive effects of agricultural landscape heterogeneity and weather conditions on breeding density and reproductive success of a diurnal raptor.

Agricultural intensification and climate change are serious threats toward animal populations worldwide. Agricultural intensification reduces the heterogeneity of agricultural habitats by diminishing crop variation and destroying microhabitats, such as small woody features, whereas the effects of climate change range from the growing frequency of weather extremes to disrupted prey-predator dynamics. We collected long-term ringing data from a population of Eurasian kestrels (Falco tinnunculus) located amidst agricultural areas in western Finland during 1985-2021, which we combined with density indices of their main prey species (voles), spatial data consisting of land cover classification of kestrel territories, and weather data, to study the effects of different environmental drivers on breeding density and success. We found that the density of inhabited nests rose with vole abundance and springtime snow depth, with the overall trend of population growth being stronger in areas with more heterogeneous landscapes. Clutch size was influenced negatively by the age of male parent and landscape heterogeneity, and positively by vole abundance, with rainfall having a negative influence conditional to other variables. Likewise, the number of produced fledglings was affected by male age, but it was additionally positively associated with landscape heterogeneity and its interaction with rainfall, with greater fledgling output in heterogeneous landscapes during high precipitation. The discrepancy between factors predicting large clutches and high numbers of fledglings suggests that while kestrels do not prefer heterogeneous landscapes when prospecting for territories, heterogeneous habitats provide better circumstances for foraging during the nestling period, which ensures nestling survival, particularly during adverse environmental conditions. Therefore, breeding in areas under intense agricultural use is more suboptimal to kestrels than their territory preferences would indicate. As changing climate may reduce prey availability and heighten the probability of weather extremities, agricultural intensification may lead to weaker reproductive success in densely populated farmland habitats.

Anthropogenic bottom-up and top-down impacts on boreal breeding waterbirds.

Wetland habitats are changing under multiple anthropogenic pressures. Nutrient leakage and pollution modify physico-chemical state of wetlands and affect the ecosystem through bottom-up processes, while alien predators affect the ecosystems in a top-down manner. Boreal wetlands are important breeding areas for several waterbird species, the abundances of which potentially reflect both bottom-up and top-down ecosystem processes. Here, we use long-term national monitoring data gathered from c. 130 waterbird breeding sites in Finland from the 1980s to the 2020s. We hypothesised that the physico-chemical state of the waters and increasing alien predator abundance both play a role in steering the waterbird population trends. We set out to test this hypothesis by relating population changes of 17 waterbird species to changes in water chemistry and to regional alien predator indices while allowing species-specific effects to vary with foraging niche (dabblers, invertivore divers, piscivorous divers, herbivores), nesting site, female mass and habitat (oligotrophic, eutrophic). We found niche and nesting site-specific, habitat-dependent changes in waterbird numbers. While the associations with higher phosphorus levels and browning water were in overall positive at the oligotrophic lakes, the numbers of invertivore and piscivore diving ducks were most strongly negatively associated with higher phosphorus levels and browning water at the eutrophic lakes. Furthermore, increased pH levels benefitted piscivores. Invertivore diving duck species nesting on the wetlands had declined most on sites with high alien predator indices. Large herbivorous species and species preferring oligotrophic lakes seem to be successful. We conclude that the large-scale breeding waterbird decline in Finland is closely connected to both bottom-up and top-down processes, where negative associations are emphasised especially at eutrophic lakes. Niche-, nest site- and habitat-specific management actions are required to conserve declining waterbird populations. Managing wetlands on catchments level together with alien predator control may provide important approaches to future wetland management.

Balancing between predation risk and food by boreal breeding ducks.

Wetlands belong to the globally most threatened habitats, and organisms depending on them are of conservation concern. Wetland destruction and quality loss may affect negatively also boreal breeding ducks in which habitat selection often needs balancing between important determinants of habitat suitability. In Finland duck population trajectories are habitat-specific, while the reasons behind are poorly understood. In this research, we studied the balance of nest predation risk and invertebrate food abundance in boreal breeding ducks in Finland at 45 lakes and ponds in 2017 and 2018. We surveyed duck pairs and broods from these and 18 additional water bodies. We evaluated nest predation by monitoring artificial nests with camera traps over a 7-day exposure period and sampled invertebrates from water bodies using emergence and activity traps. Camera trap results indicate that predation risk was higher in the water bodies surrounded by agricultural land than in forestland. Ponds (seasonal, beaver, and man-made) had lower nest predation risk, and they were also more invertebrate-rich than permanent lakes. In addition, artificial nests further away from water bodies had higher survival than shoreline nests. Habitat use of duck pairs was not associated with invertebrate food, but duck broods preferred habitats rich in food. High nest predation pressure in shorelines of especially agricultural landscapes may contribute to the declining population trends of ducks in Finland. Controlling predators could be an important conservation action to improve duck breeding success. This research underlines the benefits of the availability of different water body types for breeding ducks. There is an urgent need to pay attention to protecting seasonal ponds, while the lack of flooded waters may be mitigated by favouring beavers or creating man-made ponds.

Local variation in the timing and advancement of lake ice breakup and impacts on settling dynamics in a migratory waterbird.

Timing of ice-out is important to fundamental hydrological and ecological processes in freshwater ecosystems at high northern latitudes. While earlier ice-out in lakes during the last century is a well-documented phenomenon across the Northern Hemisphere, local variation in the rate of advancement of ice-out has received little attention. Here, records of ice-out date in 1991-2020 from 37 small lakes in a boreal catchment area in southeastern Finland were used to study variation in the timing of ice-out and its advancement. In addition, data of settling phenology of migratory common goldeneyes (Bucephala clangula) at the study lakes were used to examine how between-year and within-season variation in the timing of ice-out affects lake settlement of the species. Overall, ice-out date (IOD, the timing of ice break-up in the spring) advanced 9.8 days during the 30-year study period, April temperature being more important than winter temperature (severity) in determining the IOD. Rate of the advancement of IOD in individual lakes varied from 1.5 to 16.1 days, having advanced more in relatively larger lakes. Lakes at higher elevations had later mean IOD than lakes at lower elevations. Within-season differences among the lakes in IOD increased from 1991 to 2020, this variation being mainly driven by temperature during the ice melting period. Lakes with late mean IOD were settled later in a season by breeding common goldeneyes than lakes with early IOD. The faster the ice melting progressed within a season, the faster common goldeneyes settled the breeding lakes. The results demonstrate how global warming differently affects IOD in boreal lakes even within the same catchment area. More research in the landscape context is needed to enhance our understanding of changes in IOD in boreal lakes and how differently advancing IOD affects local dynamics of species dependent on open water.

Seasonal and diel activity patterns of the endangered taiga bean goose (Anser fabalis fabalis) during the breeding season, monitored with camera traps.

Taiga bean goose (Anser fabalis fabalis) is an endangered subspecies that breeds sporadically in remote habitats in the arctic and boreal zones. Due to its elusive behaviour, there is a paucity of knowledge on the behaviour of taiga bean goose during the breeding season, and survey methods for monitoring numbers in the breeding areas are lacking. Camera traps are a useful tool for wildlife monitoring, particularly when there is a need for non-invasive methods due to the shy nature of the species. In this study, we tested the use of camera traps to investigate seasonal and diel activity patterns of taiga bean goose in Finland over two successive breeding seasons, 2018 and 2019. We did this by modelling counts of geese from images with generalized linear and additive mixed models. The camera type (cameras placed by experts specialized in bean goose ecology vs randomly placed cameras) did not influence the count of taiga bean goose (p = 0.386). However, the activity varied significantly by region, Julian day, time of day and temperature, with the study site (individual peatland) and year adding substantial random variation and uncertainty in the counts. Altogether, the best fitting model explained nearly 70% of the variation in taiga bean goose activity. The peak in activity occurred about a month later in the southernmost region compared to the more northern regions, which may indicate behaviours related to migration rather than breeding and moulting. Our results show that long-term monitoring with game camera traps provide a potential unobtrusive approach for studying the behavioural patterns of taiga bean goose and can increase our ecological knowledge of this little-known subspecies. The results can be applied to planning of the annual censuses and finding the optimal time frame for their execution.

Plasticity in timing of avian breeding in response to spring temperature differs between early and late nesting species.

Plasticity for breeding dates may influence population vulnerability to climate change via phenological mismatch between an organism's life cycle requirements and resource availability in occupied environments. Some life history traits may constrain plasticity, however there have been remarkably few comparisons of how closely-related species, differing in key traits, respond to common phenology gradients. We compared population- and individual-level plasticity in clutch initiation dates (CID) in response to spring temperature among five duck species with early- to late-season nesting life histories. Plasticity was strongest in females of the earliest breeding species (common goldeneye [Bucephala clangula], mallard [Anas platyrhynchos], and gadwall [Mareca strepera]), whereas late-nesting lesser scaup (Aythya affinis) and white-winged scoter (Melanitta fusca deglandi) did not respond. These results contrast with previous work in other bird families that suggested late-breeders are generally more flexible. Nevertheless, late-breeding species exhibited annual variation in mean CID, suggesting response to other environmental factors unrelated to spring temperature. Goldeneye and gadwall females varied in their strength of individual plasticity ('individual × environment' interactions) and goldeneye and scoter females showed evidence of interannual repeatability of CID. Fitness consequences of CID plasticity in response to spring phenology, including trophic mechanisms and population consequences, warrant investigation.

Invertebrates are declining in boreal aquatic habitat: The effect of brownification?

Surface water browning affects boreal lakes in the Northern Hemisphere. This process is expected to increase with global warming. Boreal lakes are the most numerous lakes on Earth. These ecosystems are particularly sensitive to disturbances due to their low biodiversity compared to other aquatic environments. The recent darkening of surface water is expected to hinder key ecosystem processes, particularly through lower primary productivity and loss of biodiversity. However, studies based on long-term data collections have rarely been conducted on the ecological consequences of water browning on aquatic food webs, especially concerning its impacts on invertebrate communities. For the first time, our analysis based on two decades of data collection in Finnish lakes highlighted a relation between water browning and a decline in aquatic macroinvertebrate abundances. Aquatic invertebrates are the main food resource for many secondary predators such as fish and waterbirds, hence such effect on their populations may have major consequences for boreal ecosystem functioning.

Changes in species richness and composition of boreal waterbird communities: a comparison between two time periods 25 years apart.

Global measures of biodiversity indicate consistent decline, but trends reported for local communities are more varied. Therefore, we need better understanding of mechanisms that drive changes in diversity of local communities and of differences in temporal trends between components of local diversity, such as species richness and species turnover rate. Freshwater ecosystems are vulnerable to multiple stressors, and severe impacts on their biodiversity have been documented. We studied species richness and composition of local boreal waterbird communities in 1990/1991 and 2016 at 58 lakes distributed over six regions in Finland and Sweden. The study lakes represented not only local trophic gradients but also a latitudinal gradient in the boreal biome. While species richness tended to be lower in 2016 than in 1990/1991, species turnover was relatively high. Within foraging guilds, local species richness of diving ducks and surface feeding waterbirds decreased, whereas that of large herbivores increased. The number of species gained in local communities was higher in lakes with rich vegetation than in lakes with sparse vegetation. Conservation of boreal freshwater ecosystems would benefit from recognizing that large-scale environmental changes can affect local diversity via processes operating at finer scales.

Generalist predator, cyclic voles and cavity nests: testing the alternative prey hypothesis.

The alternative prey hypothesis (APH) states that when the density of the main prey declines, generalist predators switch to alternative prey and vice versa, meaning that predation pressure on the alternative prey should be negatively correlated with the density of the main prey. We tested the APH in a system comprising one generalist predator (pine marten, Martes martes), cyclic main prey (microtine voles, Microtus agrestis and Myodes glareolus) and alternative prey (cavity nests of common goldeneye, Bucephala clangula); pine marten is an important predator of both voles and common goldeneye nests. Specifically, we studied whether annual predation rate of real common goldeneye nests and experimental nests is negatively associated with fluctuation in the density of voles in four study areas in southern Finland in 2000-2011. Both vole density and nest predation rate varied considerably between years in all study areas. However, we did not find support for the hypothesis that vole dynamics indirectly affects predation rate of cavity nests in the way predicted by the APH. On the contrary, the probability of predation increased with vole spring abundance for both real and experimental nests. Furthermore, a crash in vole abundance from previous autumn to spring did not increase the probability of predation of real nests, although it increased that of experimental nests. We suggest that learned predation by pine marten individuals, coupled with efficient search image for cavities, overrides possible indirect positive effects of high vole density on the alternative prey in our study system.

Environmental variability and population dynamics: do European and North American ducks play by the same rules?

Density dependence, population regulation, and variability in population size are fundamental population processes, the manifestation and interrelationships of which are affected by environmental variability. However, there are surprisingly few empirical studies that distinguish the effect of environmental variability from the effects of population processes. We took advantage of a unique system, in which populations of the same duck species or close ecological counterparts live in highly variable (north American prairies) and in stable (north European lakes) environments, to distinguish the relative contributions of environmental variability (measured as between-year fluctuations in wetland numbers) and intraspecific interactions (density dependence) in driving population dynamics. We tested whether populations living in stable environments (in northern Europe) were more strongly governed by density dependence than populations living in variable environments (in North America). We also addressed whether relative population dynamical responses to environmental variability versus density corresponded to differences in life history strategies between dabbling (relatively "fast species" and governed by environmental variability) and diving (relatively "slow species" and governed by density) ducks. As expected, the variance component of population fluctuations caused by changes in breeding environments was greater in North America than in Europe. Contrary to expectations, however, populations in more stable environments were not less variable nor clearly more strongly density dependent than populations in highly variable environments. Also, contrary to expectations, populations of diving ducks were neither more stable nor stronger density dependent than populations of dabbling ducks, and the effect of environmental variability on population dynamics was greater in diving than in dabbling ducks. In general, irrespective of continent and species life history, environmental variability contributed more to variation in species abundances than did density. Our findings underscore the need for more studies on populations of the same species in different environments to verify the generality of current explanations about population dynamics and its association with species life history.

Mechanisms of density dependence in ducks: importance of space and per capita food.

The growth rate of populations usually varies over time, often in a density-dependent manner. Despite the large amount of literature on density dependence, relatively little is known of the mechanisms underlying the density-dependent processes affecting populations, especially per capita natality. We performed a 20-year study on the density dependence of brood production in two duck species differing in the stability of habitat use. Our study was conducted in a boreal watershed in southern Finland. We predicted that a diving duck common goldeneye Bucephala clangula, with more stable habitat use, would show stronger density dependence than a dabbling duck common teal Anas crecca. We investigated reproductive output in relation to the duck pair density per se as well as in relation to per capita food availability. As predicted, the reproductive output of the goldeneye showed a more density-dependent pattern than that of the teal. The number of goldeneye broods per pair decreased when the pair density increased. This was not the case with the teal. However, when the breeding success was measured by taking into account per capita food availability, both species showed density dependence. Our results imply that the occurrence of density dependent processes may vary even in sympatric ducks breeding in the same, relatively stable landscape. Our analysis also emphasizes that it is important to take into account per capita resource availability when studying the density dependence of breeding success. Both findings have important implications for the management and conservation of species.

Orthomyxo-, paramyxo- and flavivirus infections in wild waterfowl in Finland.

BACKGROUND: Screening wild birds for viral pathogens has become increasingly important. We tested a screening approach based on blood and cloacal and tracheal swabs collected by hunters to study the prevalence of influenza A, paramyxo-, flavi-, and alphaviruses in Finnish wild waterfowl, which has been previously unknown. We studied 310 blood samples and 115 mixed tracheal and cloacal swabs collected from hunted waterfowl in 2006. Samples were screened by RT-PCR and serologically by hemagglutination inhibition (HI) test or enzyme-linked immunosorbent assay (ELISA) for influenza A (FLUAV), type 1 avian paramyxo-(APMV-1), Sindbis (SINV), West Nile (WNV) and tick-borne encephalitis (TBEV) virus infections. RESULTS: FLUAV RNA was found in 13 tracheal/cloacal swabs and seven strains were isolated. Five blood samples were antibody positive. Six APMV-1 RNA-positive samples were found from which four strains were isolated, while two blood samples were antibody positive. None of the birds were positive for flavivirus RNA but three birds had flavivirus antibodies by HI test. No antibodies to SINV were detected. CONCLUSION: We conclude that circulation of both influenza A virus and avian paramyxovirus-1 in Finnish wild waterfowl was documented. The FLUAV and APMV-1 prevalences in wild waterfowl were 11.3% and 5.2% respectively, by this study. The subtype H3N8 was the only detected FLUAV subtype while APMV-1 strains clustered into two distinct lineages. Notably, antibodies to a likely mosquito-borne flavivirus were detected in three samples. The screening approach based on hunted waterfowl seemed reliable for monitoring FLUAV and APMV by RT-PCR from cloacal or tracheal samples, but antibody testing in this format seemed to be of low sensitivity.

Nest predation and the evolution of conspecific brood parasitism: from risk spreading to risk assessment.

Conspecific brood parasitism (CBP) is a taxonomically widespread reproductive tactic. One of the earliest hypotheses put forward to explain the evolution of CBP was "risk spreading"; that is, by laying eggs in more than one nest, parasites may increase the likelihood that at least one offspring will survive to independence. However, the risk spreading hypothesis, based on the assumptions of random nest predation and random selection of target nests by parasites, was theoretically refuted soon after its appearance. New results from the common goldeneye (Bucephala clangula) have revealed that nests are not predated at random and that parasites preferentially lay in safe nests. By taking into account these findings and by modifying accordingly the basic assumptions of the earlier model that refuted the risk spreading hypothesis, we built a model to address the role of nest predation in the evolution of CBP. Model simulations revealed that the selective advantage of parasitic laying, related to nest predation, is much higher than previously thought. Furthermore, the invasion probability of parasitic tactic when initially rare was reasonably high within our model framework. We show that the use of risk assessing, instead of random risk spreading, makes parasitic laying evolutionarily advantageous.

Experimental evidence for density-dependent survival in mallard (Anas platyrhynchos) ducklings.

It is unresolved to what extent waterfowl populations are regulated by density-dependent processes. By doing a 2-year crossover perturbation experiment on ten oligotrophic boreal lakes we addressed the hypothesis that breeding output is density dependent. Wing-clipped mallard (Anas platyrhynchos) hens were introduced with their own brood and then monitored for 24 days. Predicted responses were that per capita duckling and hen survival would be lower in high-density than in low-density treatments. Survival was evaluated by model fitting in program MARK. Density, year, and lake were used as main effects, while day after introduction, a weather harshness index, and presence of hens were covariates. Daily survival in ducklings was lower in the high-density treatment, but this effect was year dependent. The highest-ranking model for duckling survival also included a positive effect of duckling age and presence of hens, and a negative effect of harsh weather. Density did not affect female survival although there was a prominent year effect. The highest-ranking model for female survival also included negative effects of day after introduction and harsh weather. This is the first study to report density-dependent survival in experimentally introduced ducklings in a natural setting. Implications for population dynamics and management of harvested populations are far-reaching if such regulation occurs in some years, but not in others.

Ability to assess nest predation risk in secondary hole-nesting birds: an experimental study.

Because nest predation is the major source of nesting mortality in birds, site-specific predation risk may play an important role in determining birds' ability to select nest sites that reduce predation risk. This possibility has not been adequately tested. Here we report on 5-year experiments by which we studied, independently from birds' earlier experience with specific nest boxes, both the selection and predation risk of nest sites in the common goldeneye (Bucephala clangula). New, previously unoccupied nest boxes were erected in two habitat types on three study areas. Experimentally measured predation risk in the nest boxes varied between 0 and 1.0, i.e. goldeneye females could select a nest site along a wide gradient of possible predation-risk values. We did not find a difference in predation risk between occupied and unoccupied nest boxes, nor was the order of nest box occupation associated with predation risk. A power analysis revealed that our test had reasonably high power to reject a false null hypothesis. Our results suggest that common goldeneye females likely have not evolved an ability to assess predation risk of new, previously unoccupied nest sites.

Habitat selection rules in breeding mallards (Anas platyrhynchos): a test of two competing hypotheses.

Ideal preemption and conspecific attraction are alternative hypotheses of the habitat selection rules used by individuals. According to the former an occupied site is assumed to be preempted and therefore not available for later arriving individuals, whereas according to the latter individuals are assumed to be attracted by conspecifics to occupied sites, rather than avoiding them. We studied these competing hypotheses in breeding mallards (Anas platyrhynchos) by a cross-over experiment in 2 years, introducing birds onto lakes before migratory wild mallards arrived. If mallards use the ideal preemptive rule, breeding density of wild mallards in experimental lakes should be lower and they should be occupied less frequently than control lakes, but if mallards use the conspecific attraction rule the reverse should be true. Our results allowed us to reject the ideal preemptive rule whereas the conspecific attraction rule was to some extent supported. We discuss these findings in relation to population limitation. The results suggest that the local breeding population studied is not limited by spacing behaviour related to habitat selection.

Interspecific interactions and co-existence in dabbling ducks: observations and an experiment.

We studied the possible role of resource limitation and interspecific competition in assemblages of dabbling ducks on breeding lakes in Finland and Sweden with observational and experimental data. After initial vegetation mapping and yearly censuses of ducks in 1985-1990, we collected observational data in 1991-1994 from 28 lakes with natural populations of mallard Anas platyrhynchos and teal A. crecca. Mallard and teal co-occur over vast areas in the Holarctic and they are the only breeding dabbling ducks on many oligotrophic lakes. Both species are migratory in our study regions, teal arriving later in spring than mallards. Log-linear model analysis of observational presence/absence data revealed a positive, not a negative, association between the species. This association was independent of habitat diversity as well as of lake size. Mallard-teal interaction was also studied in a cross-over introduction experiment in 32 other lakes in two years. Wing-clipped mallards were introduced to breeding lakes before the arrival of teal to induce resource limitation and interspecific competition, hypothesized to reduce lake use by teal. The density of mallard pairs on experimental lakes was 2.9-8.0 times higher than on controls, but there was no negative response by teal to the treatment. This is the first combined observational-experimental demonstration of lack of interspecific competition in waterfowl. Our results indicate that heterospecific attraction may affect species co-existence in dabbling ducks.