Using change trajectories to study the impacts of multi-annual habitat loss on fledgling production in an old forest specialist bird.

The loss and subdivision of habitat into smaller and more spatially isolated units due to human actions has been shown to adversely affect species worldwide. We examined how changes in old forest cover during eight years were associated with the cumulative number of fledged offspring at the end of study period in Eurasian treecreepers (Certhia familiaris) in Central Finland. We were specifically interested in whether the initial level of old forest cover moderated this relation. We applied a flexible and powerful approach, latent growth curve modelling in a structural equation modeling (SEM) framework, to create trajectories describing changes in old forest cover through time, and studied how this change at both the territory core and landscape scales impacted fledging numbers. Our main finding was that at the territory core scale the negative impact of habitat loss on fledging numbers was lessened by the higher levels of initial forest cover, while no association was found at the landscape scale. Our study highlights a powerful, but currently under-utilised methodology among ecologists that can provide important information about biological responses to changes in the environment, providing a mechanistic way to study how land cover dynamics can affect species responses.

Survival of transplanted nests of the red wood ant Formica aquilonia (Hymenoptera: Formicidae): the effects of intraspecific competition and forest clear-cutting.

The fitness and survival of ant colonies depend on the resources near their nests. These resources may be limited due to poor habitat quality or by intra- and interspecific competitions, which in extreme cases may cause the ant colony to perish. We tested the effect of intraspecific competition and habitat degradation (forest clear-cutting) on colony survival by transplanting 26 nests of the red wood ant (Formica aquilonia Yarrow, 1955) in 26 different forest areas that contained 0-11 conspecific alien nests per hectare. F. aquilonia is highly dependent on canopy-dwelling aphids, thus the removal of trees should cause food limitation. During the course of the 4-year experiment, 9 of the forests were partially clear-cut. We found that while forest clear-cutting significantly decreased the colonies' survival, intraspecific competition did not. As a highly polygynous and polydomous species, F. aquilonia seems to tolerate the presence of alien conspecific colonies to a certain extent.

Habitat fragmentation and reproductive success: a structural equation modelling approach.

1. There is great interest on the effects of habitat fragmentation, whereby habitat is lost and the spatial configuration of remaining habitat patches is altered, on individual breeding performance. However, we still lack consensus of how this important process affects reproductive success, and whether its effects are mainly due to reduced fecundity or nestling survival. 2. The main reason for this may be the way that habitat fragmentation has been previously modelled. Studies have treated habitat loss and altered spatial configuration as two independent processes instead of as one hierarchical and interdependent process, and therefore have not been able to consider the relative direct and indirect effects of habitat loss and altered spatial configuration. 3. We investigated how habitat (i.e. old forest) fragmentation, caused by intense forest harvesting at the territory and landscape scales, is associated with the number of fledged offspring of an area-sensitive passerine, the Eurasian treecreeper (Certhia familiaris). We used structural equation modelling (SEM) to examine the complex hierarchical associations between habitat loss and altered spatial configuration on the number of fledged offspring, by controlling for individual condition and weather conditions during incubation. 4. Against generally held expectations, treecreeper reproductive success did not show a significant association with habitat fragmentation measured at the territory scale. Instead, our analyses suggested that an increasing amount of habitat at the landscape scale caused a significant increase in nest predation rates, leading to reduced reproductive success. This effect operated directly on nest predation rates, instead of acting indirectly through altered spatial configuration. 5. Because habitat amount and configuration are inherently strongly collinear, particularly when multiple scales are considered, our study demonstrates the usefulness of a SEM approach for hierarchical partitioning of habitat amount vs. habitat configuration in landscape ecology that may have bearing on biological conclusions.

Carotenoid composition of invertebrates consumed by two insectivorous bird species.

Dietary carotenoids are important pigments, antioxidants, and immune-stimulants for birds. Despite recent interest in carotenoids in bird ecology, we know surprisingly little about the carotenoid content of invertebrates consumed by birds. We compared carotenoid (lutein, beta-carotene, and total) concentrations in invertebrates brought to nestlings by two insectivorous passerines, the great tit, Parus major and the pied flycatcher, Ficedula hypoleuca. We also compared carotenoid levels between environments that were either polluted by heavy metals or were not polluted, because the carotenoid-based plumage color of P. major nestlings is affected by environmental pollution. Lepidopterans were the most carotenoid-rich food items and contained the largest proportion of lutein. There were no differences in carotenoid concentrations in the food items of the two bird species but P. major nestlings obtained more carotenoids from their invertebrate diet than F. hypoleuca nestlings because the P. major diet had a higher proportion of lepidopteran larvae. In polluted areas, P. major nestlings consumed lower levels of dietary carotenoids than in unpolluted areas because of temporal differences in caterpillar abundance between polluted and unpolluted sites. Our study suggests that pollution-related difference in nestling plumage color in P. major is related to varying dietary proportion of lutein-rich food items rather than pollution-related variation in insect carotenoid levels.

Fluctuating feather asymmetry in relation to corticosterone levels is sex-dependent in Eurasian treecreeper (Certhia familiaris) nestlings.

Fluctuating asymmetry (FA) has been widely used as a stress-related phenotypic marker of developmental instability. However, previous studies relating FA to various stressful conditions have produced inconsistent results and we still lack quantitative individual-level evidence that high FA is related to stress in wild vertebrate species. We studied how baseline plasma levels of corticosterone predicted FA of wing and tail feathers in free-living Eurasian treecreeper (Certhia familiaris) nestlings. We found a sex-specific association between corticosterone levels and FA: high corticosterone levels were related to an increased FA in male but not in female nestlings. These results suggest that in treecreepers, FA may correlate with individual stress hormone levels, male developmental trajectory being potentially more sensitive to stress than that of the female.

Island selection on mammalian life-histories: genetic differentiation in offspring size.

BACKGROUND: Since Darwin's pioneering work, evolutionary changes in isolated island populations of vertebrates have continued to provide the strongest evidence for the theory of natural selection. Besides macro-evolutionary changes, micro-evolutionary changes and the relative importance of natural selection vs. genetic drift are under intense investigation. Our study focuses on the genetic differentiation in morphological and life-history traits in insular populations of a small mammal the bank vole Myodes glareolus. RESULTS: Our results do not support the earlier findings for larger adult size or lower reproductive effort in insular populations of small mammals. However, the individuals living on islands produced larger offspring than individuals living on the mainland. Genetic differentiation in offspring size was further confirmed by the analyses of quantitative genetics in lab. In insular populations, genetic differentiation in offspring size simultaneously decreases the additive genetic variation (VA) for that trait. Furthermore, our analyses of differentiation in neutral marker loci (Fst) indicate that VA is less than expected on the basis of genetic drift alone, and thus, a lower VA in insular populations could be caused by natural selection. CONCLUSION: We believe that different selection pressures (e.g. higher intraspecific competition) in an insular environment might favour larger offspring size in small mammals. Island selection for larger offspring could be the preliminary mechanism in a process which could eventually lead to a smaller litter size and lower reproductive effort frequently found in insular vertebrates.

Immune defense of ants is associated with changes in habitat characteristics.

Although the immune functions of insects are known to correlate with body condition and food resources, the association between habitat structure and immune function is still largely unknown. We studied the effects of forest clear-cutting on encapsulation rate in gynes and workers in the forest-dwelling ant Formica aquilonia. Forest logging resulted in disturbed immunity in workers and gynes. Logging enhanced encapsulation reaction in gynes, whereas decreased that of workers. In gynes, there was a likely trade-off between growth and immune function that was apparent in terms of different investment in size and immune function in different habitats. In workers, however, such associations were not found. The results indicate that, because of disturbed immunity, environmental stress may increase susceptibility of wood ants to diseases and parasites in disturbed habitats.

Survival of male Tengmalm's owls increases with cover of old forest in their territory.

The loss and fragmentation of forest habitats have been considered to pose a worldwide threat to the viability of forest-dwelling animals, especially to species that occupy old forests. We investigated whether the annual survival of sedentary male Tengmalm's owls Aegolius funereus was associated with the cover of old coniferous forests in Finland. Survival and recapture probabilities varied annually with density changes in populations of the main prey (Microtus voles). When this variation was controlled for, and relationships between survival and proportions of the three different forest age classes (old-growth, middle-aged, and young) were modeled separately, the old-growth model was the most parsimonious. Survival increased with the cover of old forest, although the extent of old forest within owl territories was relatively small (mean approximately 12%, range 2-37%). This association, however, varied among years and appeared especially in years of increasing vole abundance. At such times, old forests may sustain high populations of bank voles Clethrionomys glareolus, shrews and small passerines. In addition, old forests may serve as refuges against large avian predator species, such as Ural owls Strix uralensis and goshawks Accipiter gentilis. Our results suggest that changes in habitat quality created by agriculture and forestry may have the potential to reduce adult survival, an essential component of fitness and population viability.

Heavy metal pollution disturbs immune response in wild ant populations.

Concern about the effects of environmental contaminants on immune function in both humans and wildlife is growing and practically nothing is known about this impact on terrestrial invertebrates, even though they are known to easily accumulate pollutants. We studied the effect of industrial heavy metal contamination on immune defense of a free-living wood ant (Formica aquilonia). To find out whether ants show an adapted immune function in a polluted environment, we compared encapsulation responses between local and translocated colonies. Local colonies showed higher heavy metal levels than the translocated ones but the encapsulation response was similar between the two groups, indicating that the immune system of local ants has not adapted to high contamination level. The encapsulation response was elevated in moderate whereas suppressed in high heavy metal levels suggesting higher risk for infections in heavily polluted areas.

Forest clearing and sex ratio in forest-dwelling wood ant Formica aquilonia.

Sex ratios of ants have been shown to vary with food resource levels in several studies, but it is not known whether forest clear-cutting has any effect on sex ratio of aphid-tending forest-dwelling ants. We investigated whether the offspring sex ratio of the forest dwelling ant Formica aquilonia varied as a response to clear-cutting. We found that the proportion of males was smaller in clear-cuts than in adjacent forests. Our results are among the first showing that anthropogenic changes in forest structures may have a potential to modify sex ratios of social insects and other forest-dwelling animals.

Environmental pollution has sex-dependent effects on local survival.

Environmental pollutants cause a potential hazard for survival in free-living animal populations. We modelled local survival (including emigration) by using individual mark-recapture histories of males and females in a population of a small insectivorous passerine bird, the pied flycatcher (Ficedula hypoleuca) living around a point source of heavy metals (copper smelter). Local survival of F. hypoleuca females did not differ between polluted and unpolluted environments. Males, however, showed a one-third higher local-survival probability in the polluted area. Low fledgling production was generally associated with decreased local survival, but males in the polluted area showed relatively high local survival, irrespective of their fledgling number. A possible explanation of higher local survival of males in the polluted area could be a pollution-induced change in hormone (e.g. corticosterone or testosterone) levels of males. It could make them to invest more on their own survival or affect the hormonal control of breeding dispersal. The local survival of males decreased in the polluted area over the study period along with the simultaneous decrease in heavy metal emissions. This temporal trend is in agreement with the stress hormone hypothesis.

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.

Effects of forest patch size on physiological stress and immunocompetence in an area-sensitive passerine, the Eurasian treecreeper (Certhia familiaris): an experiment.

We manipulated the primary brood size of Eurasian treecreepers (Certhia familiaris) breeding in different sized forest patches (0.5-12.8 ha) in moderately fragmented landscapes. We examined the effects of brood size manipulation (reduced, control, enlarged) and forest patch size on physiological stress (heterophil-lymphocyte ratios; H/L), body condition and cell-mediated immunocompetence (phytohaemagglutinin test). Nestlings' H/L ratios were negatively related to forest patch area in control and enlarged broods, whereas no effects were found in reduced broods. The effects of forest patch area were strongest in enlarged broods, which had, in general, twofold higher H/L ratios than control and reduced broods. The elevated H/L ratios were positively related to nestling mortality and negatively correlated with body-condition indices suggesting that the origin of stress in nestlings was mainly nutritional. Cell-mediated immunity of nestlings was not related to brood manipulation or to forest patch size. Also, the H/L ratios of adults were not related to brood manipulation or forest patch size. In addition, parental H/L ratios and body condition were not related to nestling H/L ratios. Our results suggest that during the breeding period the deleterious effects of habitat loss are seen explicitly in growing young.

Lifetime reproduction of a forest-dwelling owl increases with age and area of forests.

Loss and alteration of habitats by human actions are the largest worldwide hazard to biodiversity and viability of populations. In boreal forests of Eurasia and North America the natural habitat is changing, mainly because of forestry practices and agriculture. Although there is evidence that the diversity and abundance of animal species are lower in intensively managed than in natural forests, very little is known about how the changes in habitat composition affect reproduction and survival. The best available measure of individual performance in the wild is lifetime reproductive success (LRS), the number of offspring produced during a lifetime, because it combines both survival and reproductive success to a single measure. We show that the LRS of forest-dwelling Tengmalm's owls (Aegolius funereus) increases with the proportion of old forest in the territory because of a higher number of breeding attempts, whereas it decreases with the proportion of agricultural land because of declining fledging success in years when prey populations crashed during owl breeding. These unique results provide an interesting insight into how human influence on the landscape can affect life-history traits of animals through various pathways.

Forest fragmentation is associated with primary brood sex ratio in the treecreeper (Certhia familiaris).

We studied the primary brood sex ratio of an old-growth forest passerine, the Eurasian treecreeper (Certhia familiaris), along a gradient of forest fragmentation. We found evidence that male nestlings were more costly to produce, since they suffered twofold higher nestling mortality and were larger in body size than females. Furthermore, the proportion of males in the brood was positively associated with the provisioning rate and the amount of food delivered to the nestlings. During the first broods, a high edge density and a high proportion of pine forests around the nests were related to a decreased production of males. The densities of spiders, the main food of the treecreeper, were 38% higher on spruce trunks than on pine trunks. This suggests that pine-dominated territories with female-biased broods may have contained less food during the first broods. The observation was further supported by the fact that the feeding frequencies were lower in territories with high proportions of pines. In the second broods, territories with a high forest patch density produced female-biased broods, whereas high-quality territories with a large amount of deciduous trees and mixed forests produced male-biased broods. Our results suggest that habitat quality as measured by habitat characteristics is associated with sex allocation in free-living birds.

Forest management is associated with physiological stress in an old-growth forest passerine.

We investigated how physiological stress in an area-sensitive old-growth forest passerine, the Eurasian treecreeper (Certhia familiaris), is associated with forest fragmentation and forest structure. We found evidence that the concentrations of plasma corticosterone in chicks were higher under poor food supply in dense, young forests than in sparse, old forests. In addition, nestlings in large forest patches had lower corticosterone levels and a better body condition than in small forest patches. In general, corticosterone levels were negatively related to body condition and survival. We also found a decrease in corticosterone levels within the breeding season, which may have been a result of an increase in food supply from the first to the second broods. Our results suggest that forest fragmentation may decrease the fitness of free-living individual treecreepers.

Hatching asynchrony and brood reduction in Tengmalm's owl Aegolius funereus: the role of temporal and spatial variation in food abundance.

Hatching asynchrony is the consequence of birds initiating incubation before clutch completion. It has been suggested that variation in hatching asynchrony in owls is extensive, and therefore they should be excellent objects to study the effects of spatio-temporal variation in food abundance on this phenomenon. We examined how abundance and predictability of food affected hatching asynchrony in Tengmalm's owl Aegolius funereus (Linnaeus), which mainly feeds on voles which fluctuate in 3- to 4-year cycles in northern Europe. Hatching span averaged 6-7 days (range 0-13 days) and increased with clutch size. Food supply did not directly influence levels of hatching asynchrony but it influenced indirectly via marked among-year changes in clutch size. During the decrease phase of the vole cycle the proportion of hatchlings producing fledglings decreased with asynchrony, suggesting that chick mortality was most common among asynchronous broods when food became scarce. This finding is consistent with Lack's brood reduction hypothesis, i.e. that if food becomes scarce during the nestling period the youngest nestlings would die first without endangering the survival of the whole brood.

Survival of male Tengmalm's owls under temporally varying food conditions.

We analysed whether annual survival of male Tengmalm's owls in western Finland varies according to changes in the abundance of their main prey, Microtus voles. Our analyses were based on capture-recapture data over five 3-year population cycles of voles from 1981to 1995, each cycle consisting of consecutive years of low, increasing and decreasing vole abundance. Survival estimates of males in the increasing vole years (ca. 55%) were higher than in the decreasing vole years. In the latter case, males faced a drastic crash in Microtus vole numbers, and only about one-third of males survived over this crash. After the crash of vole populations, male survival increased rapidly (up to 76%) according to the recovery of Microtus vole populations. These results show that temporal variation in the abundance of their main prey modifies the survival of male Tengmalm's owls. In addition to survival, recapture rates also varied, largely due to the fact that in poor vole years a majority of males skipped breeding. The large among-cycle phase variation in survival (25-76%) probably creates selection for phenotypic plasticity in life-history traits related to survival and reproduction.

Prey depletion by the foraging of the Eurasian treecreeper, Certhia familiaris, on tree-trunk arthropods.

We examined to what extent breeding insectivorous treecreepers, Certhia familiaris, affect the abundance and the mean size of their prey population on the surface of tree trunks. In order to determine foraging pressure on tree trunks, we observed the parent birds' foraging behaviour in marked squares (25×25 m) at a short (10 m) and long distance (90 m) from the nest, when the nestlings were near fledging. Immediately after fledging, we measured the remaining food abundance on a sample of tree trunks near to and far from the nest. All arthropods longer than 1 mm were collected and identified to family level and their length was measured. Treecreepers foraged for significantly less time on trunks far from the nest than on trunks close to the nest (5 s vs 186 s). Consequently, treecreepers were found to deplete food abundance on trunks close to the nest. During the nesting period, parent birds removed almost twice as many spiders and other arthropods (excluding Formica ants) from the tree trunks close to the nest than from those further away. The size distribution of arthropods was the same between frequently and seldom-used tree trunks, suggesting that arthropod consumption by treecreepers was not size selective. Our direct measurements on the abundance of arthropods provide rare evidence for the ability of predators to deplete their food resources to a large extent.

Experimental increase of predation risk induces breeding dispersal of Tengmalm's owl.

Nest predation and its avoidance are critical components of an individual's fitness and play an important role in life history evolution. Almost all studies on this topic have been observational, and thus have not been able to separate the effects of individual quality, habitat selection and predation risk of given nest sites from each other. More experimental studies on nest predation and breeding dispersal, therefore, are needed to avoid confusing interpretations of the results. In western Finland, pine marten (Martes martes) predation risk was experimentally simulated at the nests of Tengmalm's owls (Aegolius funereus) by using a caged American mink (Mustela vison) as a predator. Nests without exposure to a mink served as controls. In accordance with our predictions and earlier observational studies, males exposed to simulated predation risk increased nest-hole shift and breeding dispersal distances compared to control males. Nest-hole shift and long breeding dispersal distances probably decrease the risk of nest predation, because pine martens are known to revisit nest-holes they have found.