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.

Corticosterone secretion patterns prior to spring and autumn migration differ in free-living barn swallows (Hirundo rustica L.).

Recent studies of long-distance migratory birds show that behavioural and physiological changes associated with predictable or unpredictable challenges during the annual cycle are distinctively regulated by hormones. Corticosterone is the primary energy regulating hormone in birds. Corticosterone levels are elevated during stresses but they are also modulated seasonally according to environmental conditions and life-history demands. We measured the baseline and stress-induced levels of corticosterone in the barn swallow (Hirundo rustica L.) just before spring and autumn migrations in South Africa and Finland, respectively. Barn swallows completing their pre-breeding moult had low body condition (residual body mass) and high baseline corticosterone levels in the wintering grounds. In contrast, baseline corticosterone levels in Finland were low and not related to residual mass. These data contradict the first prediction of the migration modulation hypothesis (MMH) by showing no association with baseline corticosterone levels and pre-migratory fuelling. Yet, the adrenocortical response to the capture and handling stress was notably blunted in South Africa compared to a strong response in Finland. Further, individuals that had started fuelling in Finland showed a reduced response to the handling stress. Taken together, elevated baseline corticosterone levels and high residual mass may blunt the adrenocortical response in long-distance migrants and aerial feeders such as the barn swallow. This observation lends support to the second prediction of the MMH.

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.

Seasonal variation in the regulation of redox state and some biotransformation enzyme activities in the barn swallow (Hirundo rustica L.).

Little is known of the normal seasonal variation in redox state and biotransformation activities in birds. In long-distance migratory birds, in particular, seasonal changes could be expected to occur because of the demands of migration and reproduction. In this study, we measured several redox parameters in the barn swallow (Hirundo rustica L.) during the annual cycle. We captured the wintering barn swallows before spring migration in South Africa, and we captured the barn swallows that arrived in spring, bred in summer, and migrated in autumn in Finland. The redox status and biotransformation activities of barn swallows varied seasonally. Wintering birds in South Africa had high biotransformation activities and appeared to experience oxidative stress, whereas in spring and summer, they showed relatively low redox (superoxide dismutase [SOD], catalase [CAT], and glutathione reductase [GR]) and biotransformation enzyme activities. Autumn birds had very low biotransformation enzyme activities and low indication of oxidative stress but high activity of some redox enzymes (GR and glucose 6-phosphate dehydrogenase [G6PDH]). High activities of some redox enzymes (SOD, GR, and G6PDH) seem to be related to migration, whereas low activities of some redox enzymes (SOD, CAT, and GR) may be associated with breeding. Barn swallows in South Africa may experience pollution-related oxidative stress, which may hamper interpretation of normal seasonal variation in redox parameters.

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.

Body condition is associated with adrenocortical response in the barn swallow (Hirundo rustica L.) during early stages of autumn migration.

Migration is an energy-demanding life-history period and also a significant population-limiting factor of long-distance migratory birds. It is important to understand how corticosterone, the main energy regulating hormone in birds, is associated with behavioural and physiological changes during migration. According to the migration modulation hypothesis (MMH), individual birds may express elevated levels of baseline corticosterone to facilitate fuelling, but down-regulate the adrenocortical response in order to protect skeletal muscles from the catabolic effects of the hormone. We measured the baseline and stress-induced levels of corticosterone in barn swallows (Hirundo rustica L.) during early stages of autumn migration. Here, we show that, while barn swallows clearly responded to the capture and handling stress by increasing the corticosterone level, the strength of this acute response was related to their energetic condition: birds with high body mass responded more rapidly and had lower peak values of corticosterone than lighter birds. Further, the baseline levels of corticosterone correlated negatively with the magnitude of the adrenocortical response. Barn swallows did not show elevated baseline levels of corticosterone in the course of autumn, which suggests that, instead of fuelling, the birds were actively migrating. Our results indicate that MMH also applies to aerial feeders, whose foraging habits differ from model birds of previous studies.

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.

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.

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.

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.