120-years of ecological monitoring data shows that the risk of overhunting is increased by environmental degradation for an isolated marine mammal population: The Baltic grey seal.

The Baltic Sea is home to a genetically isolated and morphologically distinct grey seal population. This population has been the subject of 120-years of careful documentation, from detailed records of bounty statistics to annual monitoring of health and abundance. It has also been exposed to a range of well-documented stressors, including hunting, pollution and climate change. To investigate the vulnerability of marine mammal populations to multiple stressors, data series relating to the Baltic grey seal population size, hunt and health were compiled, vital demographic rates were estimated, and a detailed population model was constructed. The Baltic grey seal population fell from approximately 90,000 to as few as 3000 individuals during the 1900s as the result of hunting and pollution. Subsequently, the population has recovered to approximately 55,000 individuals. Fertility levels for mature females have increased from 9% in the 1970s to 86% at present. The recovery of the population has led to demands for increased hunting, resulting in a sudden increase in annual quotas from a few hundred to 3550 in 2020. Simultaneously, environmental changes, such as warmer winters and reduced prey availability due to overfishing, are likely impacting fecundity and health. Future population development is projected for a range of hunting and environmental stress scenarios, illustrating how hunting, in combination with environmental degradation, can lead to population collapse. The current combined hunting quotas of all Baltic Nations caused a 10% population decline within three generations in 100% of simulations. To enable continued recovery of the population, combined annual quotas of less than 1900 are needed, although this quota should be re-evaluated annually as monitoring of population size and seal health continues. Sustainable management of long-lived slowly growing species requires an understanding of the drivers of population growth and the repercussions of management decisions over many decades. The case of the Baltic grey seal illustrates how long-term ecological time series are pivotal in establishing historical baselines in population abundance and demography to inform sustainable management.

Temperature synchronizes temporal variation in laying dates across European hole-nesting passerines.

Identifying the environmental drivers of variation in fitness-related traits is a central objective in ecology and evolutionary biology. Temporal fluctuations of these environmental drivers are often synchronized at large spatial scales. Yet, whether synchronous environmental conditions can generate spatial synchrony in fitness-related trait values (i.e., correlated temporal trait fluctuations across populations) is poorly understood. Using data from long-term monitored populations of blue tits (Cyanistes caeruleus, n = 31), great tits (Parus major, n = 35), and pied flycatchers (Ficedula hypoleuca, n = 20) across Europe, we assessed the influence of two local climatic variables (mean temperature and mean precipitation in February-May) on spatial synchrony in three fitness-related traits: laying date, clutch size, and fledgling number. We found a high degree of spatial synchrony in laying date but a lower degree in clutch size and fledgling number for each species. Temperature strongly influenced spatial synchrony in laying date for resident blue tits and great tits but not for migratory pied flycatchers. This is a relevant finding in the context of environmental impacts on populations because spatial synchrony in fitness-related trait values among populations may influence fluctuations in vital rates or population abundances. If environmentally induced spatial synchrony in fitness-related traits increases the spatial synchrony in vital rates or population abundances, this will ultimately increase the risk of extinction for populations and species. Assessing how environmental conditions influence spatiotemporal variation in trait values improves our mechanistic understanding of environmental impacts on populations.

Identifying the paths of climate effects on population dynamics: dynamic and multilevel structural equation model around the annual cycle.

How environmental factors influence population dynamics in long-distance migrants is complicated by the spatiotemporal diversity of the environment the individuals experience during the annual cycle. The effects of weather on several different aspects of life history have been well studied, but a better understanding is needed on how weather affects population dynamics through the different associated traits. We utilise 77 years of data from pied flycatcher (Ficedula hypoleuca), to identify the most relevant climate signals associated with population growth rate. The strongest signals on population growth were observed from climate during periods when the birds were not present in the focal location. The population decline was associated with increasing precipitation in the African non-breeding quarters in the autumn (near the arrival of migrants) and with increasing winter temperature along the migration route (before migration). The number of fledglings was associated positively with increasing winter temperature in non-breeding area and negatively with increasing winter temperature in Europe. These possible carry-over effects did not arise via timing of breeding or clutch size but the exact mechanism remains to be revealed in future studies. High population density and low fledgling production were the intrinsic factors reducing the breeding population. We conclude that weather during all seasons has the potential to affect the reproductive success or population growth rate of this species. Our results show how weather can influence the population dynamics of a migratory species through multiple pathways, even at times of the annual cycle when the birds are in a different location than the climate signal.

Adaptive responses of animals to climate change are most likely insufficient.

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

Safety in passenger ships: The influence of environmental design characteristics on people's perception of safety.

Although objective safety is a widely studied topic in ergonomics, subjective safety has received far less research attention. Nevertheless, most of human decision-making and behavior depends on how we perceive our environment. This study investigates the effects of various environmental design characteristics on people's safety perception in a passenger ship context. Five different environmental design characteristics were manipulated to increase the openness of the space or to create more clear navigation, resulting in 20 different cabin corridors for a passenger ship. Ninety-seven respondents were asked to rate these corridors on the perceived safety in an experiment. The results showed that people feel more safe when the corridors have a curved ceiling, when the walls do not have a split-level design, and when there is a view to the outside. Designers can use these insights when designing future environments.

Patterns of gene flow and selection across multiple species of Acrocephalus warblers: footprints of parallel selection on the Z chromosome.

BACKGROUND: Understanding the mechanisms and selective forces leading to adaptive radiations and origin of biodiversity is a major goal of evolutionary biology. Acrocephalus warblers are small passerines that underwent an adaptive radiation in the last approximately 10 million years that gave rise to 37 extant species, many of which still hybridize in nature. Acrocephalus warblers have served as model organisms for a wide variety of ecological and behavioral studies, yet our knowledge of mechanisms and selective forces driving their radiation is limited. Here we studied patterns of interspecific gene flow and selection across three European Acrocephalus warblers to get a first insight into mechanisms of radiation of this avian group. RESULTS: We analyzed nucleotide variation at eight nuclear loci in three hybridizing Acrocephalus species with overlapping breeding ranges in Europe. Using an isolation-with-migration model for multiple populations, we found evidence for unidirectional gene flow from A. scirpaceus to A. palustris and from A. palustris to A. dumetorum. Gene flow was higher between genetically more closely related A. scirpaceus and A. palustris than between ecologically more similar A. palustris and A. dumetorum, suggesting that gradual accumulation of intrinsic barriers rather than divergent ecological selection are more efficient in restricting interspecific gene flow in Acrocephalus warblers. Although levels of genetic differentiation between different species pairs were in general not correlated, we found signatures of apparently independent instances of positive selection at the same two Z-linked loci in multiple species. CONCLUSIONS: Our study brings the first evidence that gene flow occurred during Acrocephalus radiation and not only between sister species. Interspecific gene flow could thus be an important source of genetic variation in individual Acrocephalus species and could have accelerated adaptive evolution and speciation rate in this avian group by creating novel genetic combinations and new phenotypes. Independent instances of positive selection at the same loci in multiple species indicate an interesting possibility that the same loci might have contributed to reproductive isolation in several speciation events.

Identifying foraging habitats of Baltic ringed seals using movement data.

BACKGROUND: Identification of key foraging habitats of aquatic top predators is essential for designing effective management and conservation strategies. The Baltic ringed seal (Phoca hispida botnica) interacts with anthropogenic activities and knowledge of its spatial ecology is needed for planning population management and mitigating interactions with coastal fisheries. We investigated habitat use and foraging habitats of ringed seals (n = 26) with satellite telemetry in the northern Baltic Sea during autumn, which is important time for foraging for ringed seals. We used first passage time (FPT) approach to identify the areas of high residency corresponding to foraging areas. RESULTS: Tracked seals showed considerable movement; mean (±SD) home ranges (95 % adaptive local nearest-neighbour convex hull, a-LoCoH) were 8030 ± 4796 km(2). Two seals moved randomly and foraging areas could not be identified for them. The majority (24/26) of the studied seals occupied 1-6 main foraging areas, where they spent 47 ± 22 % of their total time. Typically the foraging areas of individuals had a mean distance of 254 ± 194 km. Most of the seals (n = 17) were "long-range foragers" which occupied several spatially remote foraging areas (mean distance 328 ± 180 km) or, in the case of two individuals, did not concentrate foraging to any particular area. The other seals (n = 9) were "local foragers" having only one foraging area or the mean distance between several areas was shorter (67 ± 26 km). Foraging areas of all seals were characterised by shallow bathymetry (median ± SD: 13 ± 49 m) and proximity to the mainland (10 ± 14 km), partly overlapping with protected areas and coastal fisheries. CONCLUSIONS: Our results indicate that in general the ringed seals range over large areas and concentrate feeding to different-often remote-areas during the open water season. Therefore, removal of individuals near the fishing gear may not be a locally effective method to mitigate seal depredation. Overlap of foraging areas with protected areas indicate that management of key foraging and resting habitats could to some extent be implemented within the existing network of marine protected areas.

A novel tool to mitigate by-catch mortality of baltic seals in coastal fyke net fishery.

Developing methods to reduce the incidental catch of non-target species is important, as by-catch mortality poses threats especially to large aquatic predators. We examined the effectiveness of a novel device, a "seal sock", in mitigating the by-catch mortality of seals in coastal fyke net fisheries in the Baltic Sea. The seal sock developed and tested in this study was a cylindrical net attached to the fyke net, allowing the seals access to the surface to breathe while trapped inside fishing gear. The number of dead and live seals caught in fyke nets without a seal sock (years 2008-2010) and with a sock (years 2011-2013) was recorded. The seals caught in fyke nets were mainly juveniles. Of ringed seals (Phoca hispida botnica) both sexes were equally represented, while of grey seals (Halichoerus grypus) the ratio was biased (71%) towards males. All the by-caught seals were dead in the fyke nets without a seal sock, whereas 70% of ringed seals and 11% of grey seals survived when the seal sock was used. The seal sock proved to be effective in reducing the by-catch mortality of ringed seals, but did not perform as well with grey seals.

Natural selection for earlier male arrival to breeding grounds through direct and indirect effects in a migratory songbird.

For migratory birds, the earlier arrival of males to breeding grounds is often expected to have fitness benefits. However, the selection differential on male arrival time has rarely been decomposed into the direct effect of male arrival and potential indirect effects through female traits. We measured the directional selection differential on male arrival time in the pied flycatcher (Ficedula hypoleuca) using data from 6 years and annual number of fledglings as the fitness proxy. Using structural equation modeling, we were able to take into account the temporal structure of the breeding cycle and the hierarchy between the examined traits. We found directional selection differentials for earlier male arrival date and earlier female laying date, as well as strong selection differential for larger clutch size. These selection differentials were due to direct selection only as indirect selection for these traits was nonsignificant. When decomposing the direct selection for earlier male arrival into direct and indirect effects, we discovered that it was almost exclusively due to the direct effect of male arrival date on fitness and not due to its indirect effects via female traits. In other words, we showed for the first time that there is a direct effect of male arrival date on fitness while accounting for those effects that are mediated by effects of the social partner. Our study thus indicates that natural selection directly favored earlier male arrival in this flycatcher population.

Selection on laying date is connected to breeding density in the pied flycatcher.

Timing of reproduction and clutch size are important determinants of breeding success, especially in seasonal environments. Several recent bird population studies have shown changes in breeding time and in natural selection on it. These changes have often been linked with climate change, but few studies have investigated how the traits or natural selection are actually connected with climatic factors. Furthermore, the effect of population density on selection has been rarely considered, despite the potential importance of density in demographic processes. We studied variation in natural selection on laying date and on clutch size in relation to measures of spring phenology and population density in a long-term study of pied flycatchers in SW Finland. The phenological stage of the environment at mean egg-laying did not affect the direction of selection on either laying date or on clutch size. There was, however, stronger selection for earlier laying date when the breeding density of the population was high, suggesting that early breeding is not necessarily beneficial as such, but that its importance is emphasized when high population density increases competition. In addition, early breeding was favoured when the pre-breeding period was cool, which may indicate an increased advantage for the fittest individuals in harsher conditions. In the middle of the twentieth century, there was selection for large clutch size, which subsequently ceased, along with an overall decrease in recruit production. Our results indicate that attention should be paid to demographic factors such as breeding density when studying natural selection and temporal changes in it.

Yolk hormones and sexual conflict over parental investment in the pied flycatcher.

Female birds might be able to manipulate the parental effort of their male partner through elevated transfer of hormones to the eggs, since these hormones affect many chick traits that males might use as cues for adjusting the level of their investment. We experimentally studied whether female pied flycatchers Ficedula hypoleuca could manipulate male investment via yolk androgens. There is much more variation in yolk androgen levels between females than within clutches, and in order to change the androgen levels of the eggs, we swapped whole clutches between nests. To estimate the androgen levels of the clutch, we measured the androgen content of a single egg per clutch. Females did not succeed in manipulating male effort using yolk androgens, since there was no relationship between the division of parental care within a pair and either original or foster egg androgen levels. One of these relationships should have occurred if females were manipulating males. The proportion of feeding visits by the male was higher when the male was old (55%) than when he was young (45%) and females laid eggs with higher androgen levels when mated with a young male. Young males did not exhibit any responses to yolk androgen levels either, which indicates that females cannot exploit their effort more than that of old males. We suggest that females may allocate yolk androgens to adjust the growth trajectories of the chicks to poor growing conditions when mated with young males that are poor providers or occupying a poor territory.

Digit ratios have poor indicator value in a wild bird population.

Early androgen exposure is known to have long-lasting effects on phenotype, behaviour and even fitness, but difficulties in measuring the exposure hinders the study of its importance in evolutionary context. Digit ratios have been highlighted as a potential easy-to-measure indicator of early steroid exposure, as they have been suggested to reflect steroid, mainly testosterone levels during prenatal development. However, evidence for digit ratios reflecting early steroid levels is weak, as experimental studies, especially in wild populations, are scarce. We studied the association between maternally derived yolk androgens and digit ratios (2D:4D, 2D:3D and 3D:4D) using both correlative data and a rather high level of experimental elevation of yolk androgens in a passerine bird, the pied flycatcher (Ficedula hypoleuca). We also examined whether digit ratios have indicator value in an evolutionary context by studying correlations between digit ratios and reproductive traits, secondary sexual traits and exploratory behaviour. We did not find any association between digit ratios and yolk androgen level either in correlative or experimental data. Digit ratios were neither related to any of the reproductive and secondary sexual traits or exploratory behaviour measured. There was, however, a sex difference in 2D:3D and 3D:4D of adult birds (due to second and fourth digits being shorter in females), which was not apparent in fledglings or captivity-raised juveniles. This suggests that either the sex difference may develop as late as during the sexual maturation for breeding. These results indicate that, in this species, digit ratios are not reliable markers of maternally derived yolk androgen exposure and that they bear little relevance as correlates of the adaptive traits we measured.

Habitat-mediated impact of alien mink predation on common frog densities in the outer archipelago of the Baltic Sea.

Alien predators have been recognised as one possible cause for amphibian declines around the world, but little is known of habitat-mediated predation impacts especially on adult amphibians. We studied common frog Rana temporaria under American mink Mustela vison predation in the outer archipelago of the Baltic Sea, south-western Finland. Using egg batches as an index of breeding frog female numbers we compared frog numbers and densities between a large, long-term mink-removal area and a comparable control area. Frog numbers in the removal area were at least 2.7-fold higher than those in the control area. In the presence of mink, frog densities increased with the amount of vegetation cover on the islands, indicating that mink predation affected frog densities especially on less-vegetated islands. An opposite trend appeared to be true for frogs in the mink-removal area, where other predators like snakes could induce a decline of frog densities on more vegetated islands. Shrub or grass vegetation seems to provide frogs shelter against alien mink predation. Our result highlights the importance of landscape-level habitat management as a conservation tool for amphibian populations.

Great tits lay increasingly smaller clutches than selected for: a study of climate- and density-related changes in reproductive traits.

1. The phenology of temperate environments and therefore timing of breeding has advanced in a number of bird species due to climate warming. Few studies, however, have examined the mechanisms behind the observed changes, the role of natural selection in them or the determinants of the selection. In other traits such as clutch size, even changes over years have been rarely studied. 2. We studied patterns and trends in timing of breeding, clutch size and fledgling production in the great tit Parus major in South-West Finland during 1953-2008, as well as natural selection on the timing and clutch size, based on fledgling production. We also examined connections between these parameters and a number of climatic and population intrinsic factors. 3. Laying date was earlier when the pre-breeding period was warm and tended to be earlier when breeding density was high, but it did not show any temporal change during the study period despite temporal increases in both explanatory factors. Number of fledglings decreased through declines in both mean clutch size and fledging success. Fledging success was better with higher breeding-time temperature and larger clutch size. Both the clutch size and fledging success were lower at higher breeding density. 4. Selection on laying date did not change through time, but there was a selection for early laying with high breeding-time temperature and high breeding density. Interestingly, in contrast to the decrease in reproductive output, the selection for larger than average clutch size strengthened with time, which was not explained by any tested factor. 5. We suggest that increasingly favourable conditions in winters have enhanced the survival and resulted in the observed increase in great tit breeding density. This may have most concerned young and otherwise low-quality individuals, which also most likely end up breeding in the increasingly occupied low-quality territories. This hypothesis was indicatively supported by increased within-year variation in both laying date and clutch size. The changes could also explain the lack of advancement in laying date as well as the increasing selection for large clutch sizes as the fittest individuals most likely occupy the best territories and lay largest clutches.

The effects of sex, age and breeding success on breeding dispersal of pied flycatchers along a pollution gradient.

We modelled breeding dispersal of an insectivorous bird, the pied flycatcher (Ficedula hypoleuca) around a point source of heavy metals (a copper smelter). We tested for the effects of sex, age, breeding success and environmental pollution on breeding dispersal distances of F. hypoleuca females and males. Unlike many earlier studies on breeding dispersal, we took into account distance-dependent sampling bias by including in our model the recapture probabilities at different distances from the site of origin. Our results show that F. hypoleuca females disperse much farther (on average 7.9 km) from their breeding sites than what was previously thought. In contrast, males only disperse short distances (on average 190 m). Breeding success affected female breeding dispersal distances depending on female age: young females moved on average 8 km from their previous breeding place irrespective of their breeding success, while old females only seemed to move this far when their fledgling production was good. Females successful in their breeding dispersed as far as less successful females, or, among old birds, even farther. Females which dispersed long distances produced more fledglings after the movement than those staying near their previous breeding site. Degree of environmental pollution had no effect on female or male breeding dispersal distances. A polluted and unproductive environment does not seem to stimulate F. hypoleuca parents to move to more profitable territories.

Climate change can alter competitive relationships between resident and migratory birds.

Climate change could affect resource competition between resident and migratory bird species by changing the interval between their onsets of breeding or by altering their population densities. We studied interspecific nest-hole competition between resident great tits and migrant pied flycatchers in South-Western Finland over the past five decades (1953-2005). We found that appearance of fatal take-over trials, the cases where a pied flycatcher tried to take over a great tit nest but was killed by the tit, increased with a reduced interspecific laying date interval and with increasing densities of both tits and flycatchers. The probability of pied flycatchers taking over great tit nests increased with the density of pied flycatchers. Laying dates of the great tit and pied flycatcher are affected by the temperatures of different time periods, and divergent changes in these temperatures could consequently modify their competitive interactions. Densities are a result of reproductive success and survival, which can be affected by separate climatic factors in the resident great tit and trans-Saharan migrant pied flycatcher. On these bases we conclude that climate change has a great potential to alter the competitive balance between these two species.

Alien mink predation induces prolonged declines in archipelago amphibians.

Amphibians are undergoing enigmatic global declines variously attributed to a complex web of anthropogenic forces. Alien predators pose a fundamental threat to biodiversity generally that is predicted to be most acute in island ecosystems. While amphibian eggs and tadpoles are vulnerable to aquatic predators, the effect of predators on adult, reproducing frogs, which most influence amphibian population processes, is unknown. Here, we report on the responses of amphibian populations in the outer Finnish Archipelago to a long-term, large-scale removal of American mink (Mustela vison Schreb.), an invasive predator linked to recent biodiversity loss across Europe. Removal increased both the densities and distribution of common frogs (Rana temporaria L.) but not those of common toads (Bufo bufo L.), which appear to escape mink predation because of their unpalatable skin. Importantly, the largest benefits of mink removal to frog recovery were slow to appear as frogs apparently have a delayed maturation in these harsh environments, which means we must be cautious about reliance upon short-term results.