Maternally transferred thyroid hormones and life-history variation in birds.

In vertebrates, thyroid hormones (THs) play an important role in the regulation of growth, development, metabolism, photoperiodic responses and migration. Maternally transferred THs are important for normal early phase embryonic development when embryos are not able to produce endogenous THs. Previous studies have shown that variation in maternal THs within the physiological range can influence offspring phenotype. Given the essential functions of maternal THs in development and metabolism, THs may be a mediator of life-history variation across species. We tested the hypothesis that differences in life histories are associated with differences in maternal TH transfer across species. Using birds as a model, we specifically tested whether maternally transferred yolk THs covary with migratory status, developmental mode and traits related to pace-of-life (e.g. basal metabolic rate, maximum life span). We collected un-incubated eggs (n = 1-21 eggs per species, median = 7) from 34 wild and captive bird species across 17 families and six orders to measure yolk THs [both triiodothyronine (T3) and thyroxine (T4)], compiled life-history trait data from the literature and used Bayesian phylogenetic mixed models to test our hypotheses. Our models indicated that both concentrations and total amounts of the two main forms of THs (T3 and T4) were higher in the eggs of migratory species compared to resident species, and total amounts were higher in the eggs of precocial species, which have longer prenatal developmental periods, than in those of altricial species. However, maternal yolk THs did not show clear associations with pace-of-life-related traits, such as fecundity, basal metabolic rate or maximum life span. We quantified interspecific variation in maternal yolk THs in birds, and our findings suggest higher maternal TH transfer is associated with the precocial mode of development and migratory status. Whether maternal THs represent a part of the mechanism underlying the evolution of precocial development and migration or a consequence of such life histories is currently unclear. We therefore encourage further studies to explore the physiological mechanisms and evolutionary processes underlying these patterns.

Northward expanding resident species benefit from warming winters through increased foraging rates and predator vigilance.

Species distributions shift northwards due to climate change, but the ecological mechanisms allowing range expansions are not fully understood. Most studies have concentrated on breeding seasons, but winter warming may also be important. Wintering distributions are restricted by food availability and temperature, which may also interact. Foraging in cold conditions requires adaptations as individuals have to be efficient in foraging, while staying warm and vigilant for predators. When the ambient temperature declines, foraging rates should be reduced due to increased time spent on warming behaviours. In addition, predator vigilance should decline, because more time has to be invested in foraging. Cold weather should limit northward expanding southern species in particular, while northern species should perform better in cold conditions. We tested this by studying temperature responses (between 0 and - 35 °C) among wintering birds at feeders. We compared foraging behaviours of two northward expanding southern species, the great tit (Parus major) and the blue tit (Cyanistes caeruleus) to a northern species, the willow tit (Poecile montanus). Foraging rate and vigilance decreased, and warming behaviour increased when temperatures declined. Importantly, the performance in these traits was poorer in the southern species compared to the willow tit. Furthermore, the response to decreasing temperatures in foraging rates and warming behaviour was stronger in the great tits than willow tits. As the winters become warmer, these mechanisms should increase wintering success of southern species wintering at high latitudes, and lead to higher survival, increased population growth, and consequent range expansion.

Larval melanism in a geometrid moth: promoted neither by a thermal nor seasonal adaptation but desiccating environments.

Spatiotemporal variation in the degree of melanism is often considered in the context of thermal adaptation, melanism being advantageous under suboptimal thermal conditions. Yet, other mutually nonexclusive explanations exist. Analysis of geographical patterns combined with laboratory experiments on the mechanisms of morph induction helps to unveil the adaptive value of particular cases of polyphenism. In the context of the thermal melanism hypothesis and seasonal adaptations, we explored an array of environmental factors that may affect the expression and performance of nonmelanic vs. melanic larval morphs in different latitudinal populations of the facultatively bivoltine moth Chiasmia clathrata (Lepidoptera: Geometridae). Geographical variation in larval coloration was independent of average temperatures experienced by the populations in the wild. The melanic morph was, however, more abundant in dry than in mesic habitats. In the laboratory, the melanic morph was induced especially under a high level of incident radiation but also at relatively high temperatures, but independently of photoperiod. Melanic larvae had higher growth rates and shorter development times than the nonmelanic ones when both temperature and the level of incident radiation were high. Our results that melanism is induced and advantageous in warm desiccating conditions contradict the thermal melanism hypothesis for this species. Neither has melanism evolved to compensate time constraints due to forthcoming autumn. Instead, larvae solve seasonal variation in the time available for growth by an elevated growth rate and a shortened larval period in the face of autumnal photoperiods. The phenotypic response to the level of incident radiation and a lack of adaptive adjustment of larval growth trajectories in univoltine populations underpin the role of deterministic environmental variation in the evolution of irreversible adaptive plasticity and seasonal polyphenism.

Informed renesting decisions: the effect of nest predation risk.

Animals should cue on information that predicts reproductive success. After failure of an initial reproductive attempt, decisions on whether or not to initiate a second reproductive attempt may be affected by individual experience and social information. If the prospects of breeding success are poor, long-lived animals in particular should not invest in current reproductive success (CRS) in case it generates costs to future reproductive success (FRS). In birds, predation risk experienced during breeding may provide a cue for renesting success. Species having a high FRS potential should be flexible and take predation risk into account in their renesting decisions. We tested this prediction using breeding data of a long-lived wader, the southern dunlin Calidris alpina schinzii. As predicted, dunlin cued on predation risk information acquired from direct experience of nest failure due to predation and ambient nest predation risk. While the overall renesting rate was low (34.5%), the early season renesting rate was high but declined with season, indicating probable temporal changes in the costs and benefits of renesting. We develop a conceptual cost-benefit model to describe the effects of the phase and the length of breeding season on predation risk responses in renesting. We suggest that species investing in FRS should not continue breeding in short breeding seasons in response to predation risk but without time constraints, their response should be similar to species investing in CRS, e.g. within-season dispersal and increased nest concealment.

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

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

Survival fluctuation is linked to precipitation variation during staging in a migratory shorebird.

Understanding how weather conditions affect animal populations is essential to foresee population changes in times of global climate shifts. However, assessing year-round weather impacts on demographic parameters is hampered in migratory animals due to often unknown occurrence in space and time. We addressed this by coupling tracking and weather data to explain extensive variation in apparent survival across 19 years in a northern European population of little ringed plovers (Charadrius dubius). Over 90% (n = 21) of tracked individuals followed migration routes along the Indo-European flyway to south India. Building on capture-recapture histories of nearly 1400 individuals, we found that between-year variation in precipitation during post-breeding staging in northern South Asia explained 47% of variation in apparent adult survival. Overall, the intensity of the monsoon in South Asia explained 31-33% of variability in apparent survival. In contrast, weather conditions in breeding, final non-breeding and pre-breeding quarters appeared less important in this species. The integration of multi-source data seems essential for identifying key regions and periods limiting population growth, for forecasting future changes and targeting conservation efforts.

Absence of Francisella tularensis in Finnish Ixodes ricinus and Ixodes persulcatus ticks.

Francisella tularensis subsp. holarctica is the causative agent of tularaemia in Europe. Finland is a high-incidence region for tularaemia, with mosquito bites as the most common sources of infection. However, in Central and Western Europe, ticks (Acari: Ixodidae) have been suggested as the main vectors. Indeed, several studies have reported the pathogen from the locally most common human-biting tick species, Ixodes ricinus. In Finland, the occurrence of the pathogen in ticks has started receiving attention only recently. Here, we collate previous tick screening data from Finland regarding F. tularensis as well as present the results from a novel screening of roughly 15 000 I. ricinus and I. persulcatus collected from across the country. In total, 14 878 ticks collected between 2015 and 2020 were screened for F. tularensis using a TaqMan-based qPCR assay targeting the 23 KDa gene. The combined screening efforts of the current and previous studies, encompassing roughly 20 000 ticks, did not find any positive ticks. Given the negative results despite the considerable sample size, it appears that the pathogen is not circulating in local tick populations in Finland. We discuss some possible reasons for the lack of the bacterium in ticks in this high-incidence region of tularaemia.

Genetic differentiation in an endangered and strongly philopatric, migrant shorebird.

BACKGROUND: Populations living in fragmented habitats may suffer from loss of genetic variation and reduced between-patch dispersal, which are processes that can result in genetic differentiation. This occurs frequently in species with reduced mobility, whereas genetic differentiation is less common among mobile species such as migratory birds. The high dispersal capacity in the latter species usually allows for gene flow even in fragmented landscapes. However, strongly philopatric behaviour can reinforce relative isolation and the degree of genetic differentiation. The Southern Dunlin (Calidris alpina schinzii) is a philopatric, long-distance migratory shorebird and shows reduced dispersal between isolated breeding patches. The endangered population of the Southern Dunlin breeding at the Baltic Sea has suffered from habitat deterioration and fragmentation of coastal meadows. We sampled DNA across the entire population and used 12 polymorphic microsatellite loci to examine whether the environmental changes have resulted in genetic structuring and loss of variation. RESULTS: We found a pattern of isolation-by-distance across the whole Baltic population and genetic differentiation between local populations, even within the southern Baltic. Observed heterozygosity was lower than expected throughout the range and internal relatedness values were positive indicating inbreeding. CONCLUSIONS: Our results provide long-term, empirical evidence for the theoretically expected links between habitat fragmentation, population subdivision, and gene flow. They also demonstrate a rare case of genetic differentiation between populations of a long-distance migratory species. The Baltic Southern Dunlin differs from many related shorebird species that show near panmixia, reflecting its philopatric life history and the reduced connectivity of its breeding patches. The results have important implications as they suggest that reduced connectivity of breeding habitats can threaten even long-distance migrants if they show strong philopatry during breeding. The Baltic Southern Dunlin warrants urgent conservation efforts that increase functional connectivity and gene flow between breeding areas.

Questing abundance of adult taiga ticks Ixodes persulcatus and their Borrelia prevalence at the north-western part of their distribution.

BACKGROUND: Because ixodid ticks are vectors of zoonotic pathogens, including Borrelia, information of their abundance, seasonal variation in questing behaviour and pathogen prevalence is important for human health. As ticks are invading new areas northwards, information from these new areas are needed. Taiga tick (Ixodes persulcatus) populations have been recently found at Bothnian Bay, Finland. We assessed seasonal variation in questing abundance of ticks and their pathogen prevalence in coastal deciduous forests near the city of Oulu (latitudes 64-65°) in 2019. METHODS: We sampled ticks from May until September by cloth dragging 100 meters once a month at eight study sites. We calculated a density index (individuals/100 m2) to assess seasonal variation. Samples were screened for Borrelia burgdorferi (sensu lato) (including B. afzelii, B. garinii, B. burgdorferi (sensu stricto) and B. valaisana), Borrelia miyamotoi, Anaplasma phagocytophilum, Rickettsia spp., Neoehrlichia mikurensis, Francisella tularensis and Bartonella spp., Babesia spp. and for the tick-borne encephalitis virus. RESULTS: All except one nymph were identified as I. persulcatus. The number of questing adults showed a strong peak in May (median: 6.5 adults/100 m2), which is among the highest values reported in northern Europe, and potentially indicates a large population size. After May, the number of questing adults declined steadily with few adults still sampled in August. Nymphs were present from May until September. We found a striking prevalence of Borrelia spp. in adults (62%) and nymphs (40%), with B. garinii (51%) and B. afzelii (63%) being the most common species. In addition, we found that 26% of infected adults were coinfected with at least two Borrelia genospecies, mainly B. garinii and B. afzelii, which are associated with different host species. CONCLUSIONS: The coastal forest environments at Bothnian Bay seem to provide favourable environments for I. persulcatus and the spread of Borrelia. High tick abundance, a low diversity of the host community and similar host use among larvae and nymphs likely explain the high Borrelia prevalence and coinfection rate. Research on the infestation of the hosts that quantifies the temporal dynamics of immature life stages would reveal important aspects of pathogen circulation in these tick populations.

Quantitative genetics of the use of conspecific and heterospecific social cues for breeding site choice.

Social information use for decision-making is common and affects ecological and evolutionary processes, including social aggregation, species coexistence, and cultural evolution. Despite increasing ecological knowledge on social information use, very little is known about its genetic basis and therefore its evolutionary potential. Genetic variation in a trait affecting an individual's social and nonsocial environment may have important implications for population dynamics, interspecific interactions, and, for expression of other, environmentally plastic traits. We estimated repeatability, additive genetic variance, and heritability of the use of conspecific and heterospecific social cues (abundance and breeding success) for breeding site choice in a population of wild collared flycatchers Ficedula albicollis. Repeatability was found for two social cues: previous year conspecific breeding success and previous year heterospecific abundance. Yet, additive genetic variances for these two social cues, and thus heritabilities, were low. This suggests that most of the phenotypic variation in the use of social cues and resulting conspecific and heterospecific social environment experienced by individuals in this population stems from phenotypic plasticity. Given the important role of social information use on ecological and evolutionary processes, more studies on genetic versus environmental determinism of social information use are needed.

Grazed wet meadows are sink habitats for the southern dunlin (Calidris alpina schinzii) due to nest trampling by cattle.

The effect of habitat management is commonly evaluated by measuring population growth, which does not distinguish changes in reproductive success from changes in survival or the effects of immigration or emigration. Management has rarely been evaluated considering complete life cycle of the target organisms, including also possible negative impacts from management. We evaluated the effectiveness of cattle grazing in the restoration of coastal meadows as a breeding habitat for small and medium-sized ground-nesting birds by examining the size and demography of a southern dunlin (Calidris alpina schinzii) breeding population. Using a stochastic renesting model that includes within-season variation in breeding parameters, we evaluated the effect of grazing time and stocking rates on reproduction. The census data indicated that the population was stable when nest trampling was prevented, but detailed demographic models showed that the population on managed meadows was a sink that persisted by attracting immigrants. Even small reductions in reproductive success caused by trampling were detrimental to long-term viability. We suggest that the best management strategy is to postpone grazing to after the 19th of June, which is about three weeks later than what is optimal from the farmer's point of view. The differing results from the two evaluation approaches warn against planning and evaluating management only based on census population size and highlight the need to consider target-specific life history characteristics and demography. Even though grazing management is crucial for creating and maintaining suitable habitats, we found that it was insufficient in maintaining a viable population without additional measures that increase nest success. In the presently studied case and in populations with similar breeding cycles, impacts from nest trampling can be avoided by starting grazing when about 70% of the breeding season has past.

Different Ultimate Factors Define Timing of Breeding in Two Related Species.

Correct reproductive timing is crucial for fitness. Breeding phenology even in similar species can differ due to different selective pressures on the timing of reproduction. These selection pressures define species' responses to warming springs. The temporal match-mismatch hypothesis suggests that timing of breeding in animals is selected to match with food availability (synchrony). Alternatively, time-dependent breeding success (the date hypothesis) can result from other seasonally deteriorating ecological conditions such as intra- or interspecific competition or predation. We studied the effects of two ultimate factors on the timing of breeding, synchrony and other time-dependent factors (time-dependence), in sympatric populations of two related forest-dwelling passerine species, the great tit (Parus major) and the willow tit (Poecile montanus) by modelling recruitment with long-term capture-recapture data. We hypothesized that these two factors have different relevance for fitness in these species. We found that local recruitment in both species showed quadratic relationships with both time-dependence and synchrony. However, the importance of these factors was markedly different between the studied species. Caterpillar food played a predominant role in predicting the timing of breeding of the great tit. In contrast, for the willow tit time-dependence modelled as timing in relation to conspecifics was more important for local recruitment than synchrony. High caterpillar biomass experienced during the pre- and post-fledging periods increased local recruitment of both species. These contrasting results confirm that these species experience different selective pressures upon the timing of breeding, and hence responses to climate change may differ. Detailed information about life-history strategies is required to understand the effects of climate change, even in closely related taxa. The temporal match-mismatch hypothesis should be extended to consider subsequent critical periods when food needs to be abundantly available.

Effects of geolocators on hatching success, return rates, breeding movements, and change in body mass in 16 species of Arctic-breeding shorebirds.

BACKGROUND: Geolocators are useful for tracking movements of long-distance migrants, but potential negative effects on birds have not been well studied. We tested for effects of geolocators (0.8-2.0 g total, representing 0.1-3.9 % of mean body mass) on 16 species of migratory shorebirds, including five species with 2-4 subspecies each for a total of 23 study taxa. Study species spanned a range of body sizes (26-1091 g) and eight genera, and were tagged at 23 breeding and eight nonbreeding sites. We compared breeding performance and return rates of birds with geolocators to control groups while controlling for potential confounding variables. RESULTS: We detected negative effects of tags for three small-bodied species. Geolocators reduced annual return rates for two of 23 taxa: by 63 % for semipalmated sandpipers and by 43 % for the arcticola subspecies of dunlin. High resighting effort for geolocator birds could have masked additional negative effects. Geolocators were more likely to negatively affect return rates if the total mass of geolocators and color markers was 2.5-5.8 % of body mass than if tags were 0.3-2.3 % of body mass. Carrying a geolocator reduced nest success by 42 % for semipalmated sandpipers and tripled the probability of partial clutch failure in semipalmated and western sandpipers. Geolocators mounted perpendicular to the leg on a flag had stronger negative effects on nest success than geolocators mounted parallel to the leg on a band. However, parallel-band geolocators were more likely to reduce return rates and cause injuries to the leg. No effects of geolocators were found on breeding movements or changes in body mass. Among-site variation in geolocator effect size was high, suggesting that local factors were important. CONCLUSIONS: Negative effects of geolocators occurred only for three of the smallest species in our dataset, but were substantial when present. Future studies could mitigate impacts of tags by reducing protruding parts and minimizing use of additional markers. Investigators could maximize recovery of tags by strategically deploying geolocators on males, previously marked individuals, and successful breeders, though targeting subsets of a population could bias the resulting migratory movement data in some species.