Effects of forest management on the spatial distribution of the willow tit (Poecile montanus)

Modification, fragmentation and loss of boreal forest habitats have been intensive during the last century due to forestry practises and land use. This has been related to population declines of many forest species, yet the mechanisms affecting on the background are largely unknown. The willow tit, a primary cavity-nesting species that was once the 4th most common bird species in Finland is nowadays endangered. Earlier findings suggest that the willow tit population is affected by the reduction of nesting sites, decaying snags in forests and the loss of mature forests which contain the food storages during the winter. In this study we are searching for the mechanisms how the forest management methods could explain the decline of the willow tit population. We used long-term breeding data of the willow tit nesting sites from 1990 to 2020 collected in a study area in northern Finland to analyse if forest management affected nearest neighbour distances and natal dispersal and breeding dispersal distances. We used Geographic Information System (GIS) methods to combine the ecological breeding data to accurate spatial forest management and habitat quality data. The data was analysed with linear mixed models. We found that clear-cuttings affected the willow tit dispersal and neighbouring nest distances more than thinnings. Both clear-cuttings and thinnings increased the nearest neighbour distances. The natal and breeding dispersal distances were lengthened by increasing proportions of clear-cuttings. The habitat loss caused by clear-cuttings and the decrease in habitat quality caused by thinnings has had a major role in the decline of the willow tit population. The forest management actions were estimated to explain about 65 % of the willow tit breeding density decrease in the study area. The effects of forest management were witnessed in a cumulative 0-30-year period meaning that forest management causes long-term habitat degradation and loss. Availability of deciduous snags in the forests can compensate the habitat loss to some extent by providing better breeding opportunities. As the effects of clear-cutting were more severe to the willow tit than thinning, we recommend using other forest management methods than clear-cutting as the main management method.Peer reviewe

From feces to data : A metabarcoding method for analyzing consumed and available prey in a bird-insect food web

Diets play a key role in understanding trophic interactions. Knowing the actual structure of food webs contributes greatly to our understanding of biodiversity and ecosystem functioning. The research of prey preferences of different predators requires knowledge not only of the prey consumed, but also of what is available. In this study, we applied DNA metabarcoding to analyze the diet of 4 bird species (willow tits Poecile montanus, Siberian tits Poecile cinctus, great tits Parus major and blue tits Cyanistes caeruleus) by using the feces of nestlings. The availability of their assumed prey (Lepidoptera) was determined from feces of larvae (frass) collected from the main foraging habitat, birch (Betula spp.) canopy. We identified 53 prey species from the nestling feces, of which 11 (21%) were also detected from the frass samples (eight lepidopterans). Approximately 80% of identified prey species in the nestling feces represented lepidopterans, which is in line with the earlier studies on the parids' diet. A subsequent laboratory experiment showed a threshold for fecal sample size and the barcoding success, suggesting that the smallest frass samples do not contain enough larval DNA to be detected by high-throughput sequencing. To summarize, we apply metabarcoding for the first time in a combined approach to identify available prey (through frass) and consumed prey (via nestling feces), expanding the scope and precision for future dietary studies on insectivorous birds.Peer reviewe

Bird populations most exposed to climate change are less sensitive to climatic variation

The phenology of many species shows strong sensitivity to climate change; however, with few large scale intra-specific studies it is unclear how such sensitivity varies over a species' range. We document large intra-specific variation in phenological sensitivity to temperature using laying date information from 67 populations of two co-familial European songbirds, the great tit (Parus major) and blue tit (Cyanistes caeruleus), covering a large part of their breeding range. Populations inhabiting deciduous habitats showed stronger phenological sensitivity than those in evergreen and mixed habitats. However, populations with higher sensitivity tended to have experienced less rapid change in climate over the past decades, such that populations with high phenological sensitivity will not necessarily exhibit the strongest phenological advancement. Our results show that to effectively assess the impact of climate change on phenology across a species' range it will be necessary to account for intra-specific variation in phenological sensitivity, climate change exposure, and the ecological characteristics of a population. Intra-specific variations may contribute to heterogeneous responses to climate change across a species' range. Here, the authors investigate the phenology of two bird species across their breeding ranges, and find that their sensitivity to temperature is uncoupled from exposure to climate change.Peer reviewe

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

Publisher Copyright: © 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.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.Peer reviewe

Boreal populations facing climatic and habitat changes

Abstract Anthropogenic climate change and habitat loss and deterioration affect populations worldwide. Climate warming has changed phenologies of many species across trophic levels. Some predator populations now experience temporal mismatches with their prey, as timings of peak prey abundance and of the predator’s highest food demands no longer meet. The temporal mismatch hypothesis suggests that the population’s recruitment rate is related to its degree of synchrony with the food resources needed to feed offspring. However, species’ and populations’ responses to climate warming differ. Human land use alters and destroys habitats of countless species. For example, many boreal forest bird populations have declined, presumably due to intensive forestry. It has decreased the amount of dead wood, causing a threat to saproxylic species. Identification of the key characteristics of high-quality habitats is essential for conservation planning and for developing sustainable forestry. As individuals are suspected to settle in habitats that maximize their fitness, analysis of nest site selection can be used to identify the key habitats. My dissertation concerns the impacts of climate change and habitat deterioration on boreal populations. I use hole-nesting passerines as model species. By utilizing long-term data I show that breeding phenologies of Parus major and Poecile montanus, but not of Cyanistes caeruleus, have shifted earlier. Also, the timing of the food peak has advanced, improving the synchrony between P. montanus and caterpillars. In P. major and C. caeruleus, synchrony has remained good. However, the positive effect of good synchrony on breeding success seems to be conditional, arising only in certain circumstances, such as in years of high caterpillar abundance. I suggest that in boreal populations temperature affects timing of breeding mostly as a proximate factor. The availability of standing decaying trees used for nesting sites was the most important habitat characteristic determining the nest site selection of P. montanus. Remote sensing data alone was insufficient to produce reliable models, as the ecologically important small-scale factor can only be determined by direct field surveys. Omission of forest thinning in selected forest sites would ensure the continuous availability of decaying wood with positive influence on biodiversity in managed forests.Tiivistelmä Ihmisen aiheuttama habitaattien katoaminen ja huononeminen sekä ilmastonmuutos vaikuttavat populaatioihin kaikkialla maailmassa. Ilmaston lämpeneminen on muuttanut monien lajien fenologioita eri trofiatasoilla. Osalla saalistajalajien populaatioista suurin ravinnontarve ei enää ajoitu samaan aikaan saaliin runsaushuipun kanssa. Ajoituksen eriaikaisuus -hypoteesin mukaan todennäköisyys populaatioon rekrytoitumiselle riippuu synkronian asteesta saaliin kanssa. Ilmaston lämpenemisen vaikutuksissa lajeihin ja populaatioihin on kuitenkin vaihtelua. Ihmisen maankäyttö muuttaa ja tuhoaa lukuisien lajien elinympäristöjä. Esimerkiksi useiden boreaalisten metsien lintupopulaatioiden pienentymistä on selitetty intensiivisellä metsätaloudella. Lahopuun määrä metsissä on vähentynyt, mikä on uhka lahopuusta riippuvaisille lajeille. Korkealaatuisten habitaattien keskeisten piirteiden tunnistaminen on tärkeää luonnonsuojelun ja kestävän metsätalouden suunnittelulle. Koska yksilöiden oletetaan valitsevan niiden kelpoisuutta maksimoivia elinympäristöjä, pesäpaikanvalinta-analyysiä voidaan käyttää tärkeiden habitaattipiirteiden tunnistamiseen. Tarkastelen väitöskirjassani ilmastonmuutoksen ja habitaattien laadun vaikutuksia boreaalisiin populaatioihin. Mallilajeina käytän koloissa pesiviä varpuslintuja. Hyödyntämällä pitkäaikaisaineistoja osoitan, että lisääntymisen ajoittuminen on aikaistunut tali- ja hömötiaisella, mutta ei sinitiaisella. Myös ravintohuippu on aikaistunut, mikä on parantanut synkroniaa hömötiaisen ja sen pääasiallisen ravinnon eli toukkien välillä. Tali- ja sinitiaisella synkronia on pysynyt hyvänä. Hyvän synkronian myönteinen vaikutus lisääntymismenestykseen vaikuttaa kuitenkin ehdolliselta: se tulee esiin vain tietyissä olosuhteissa, kuten vuosina jolloin toukkia on runsaasti. Kevään lämpötilat näyttävät vaikuttavan pesinnän ajoittumiseen erityisesti proksimaattisena tekijänä. Pesäpaikkoina toimivien seisovien lahopuiden määrä on tärkein hömötiaisen pesäpaikanvalintaa määräävä tekijä. Kaukokartoitusaineisto yksinään ei riitä luotettavien mallien tuottamiseen, sillä ekologisesti tärkeät pienen skaalan tekijät voidaan kartoittaa vain suorin maastomittauksin. Metsien harventamatta jättäminen valituilla laikuilla turvaisi lahopuun jatkuvan saatavuuden, mikä vaikuttaisi myönteisesti talousmetsien biodiversiteettiin

Effects of ambient temperatures on evolutionary potential of reproductive timing in boreal passerines

Many populations need to adapt to changing environmental conditions, such as warming climate. Changing conditions generate directional selection for traits critical for fitness. For evolutionary responses to occur, these traits need to be heritable. However, changes in environmental conditions can alter the amount of heritable variation a population expresses, making predictions about expected responses difficult. The aim of this study was to evaluate the effects of ambient temperatures on evolutionary potential and strength of natural selection on the timing of reproduction in two passerine birds breeding in boreal forests. Long-term data on individually marked Willow Tits Poecile montanus (1975-2018) and Great Tits Parus major (1969-2018) were analysed with random regression animal models to assess if spring temperatures affect the expressed amount of additive genetic variation (V-A) and heritability (h(2)) in the timing of breeding. We assessed if ambient temperatures of different seasons influenced the direction and strength of selection on breeding time. We also evaluated if the strength of selection covaried with evolutionary potential. Levels of V-A or h(2) expressed in laying date were unaffected by spring temperatures in both study species. Selection for earlier breeding was found in the Willow Tit, but not in the Great Tit. In the Willow Tit, selection for earlier breeding was more intense when the temperatures of following autumns and winters were low. Different measures of evolutionary potential did not covary strongly with the strength of selection in either species. We conclude that there is no or little evidence that climate warming would either constrain or promote evolutionary potential in timing of breeding through changes in amount of genetic variance expressed in boreal Willow and Great Tits. However, selection on the timing of breeding, a life-history event taking place in springtime, is regulated by temperatures of autumns and winters. Rapid warming of these periods have thus potential to reduce the rate of expected evolutionary response in reproductive timing.Peer reviewe

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

TIT DATA_PAKANEN ET AL

Capture-recapture data with individual covariates

Data from: 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