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The potential for climate change to disrupt phenology‐mediated interactions in interaction networks has attracted considerable attention in recent decades. Frequently, studies emphasize the fragility of ephemeral seasonal interactions, and the risks posed by phenological asynchrony. Here, we argue that the fitness consequences of asynchrony in phenological interactions may often be more buffered than is typically acknowledged. We identify three main forms that buffering may take: (i) mechanisms that reduce asynchrony between consumer and resource; (ii) mechanisms that reduce the costs of being asynchronous; and (iii) mechanisms that dampen interannual variance in performance across higher organizational units. Using synchrony between the hatching of winter moth caterpillars and the leafing of their host‐plants as a case study, we identify a wide variety of buffers that reduce the detrimental consequences of phenological asynchrony on caterpillar individuals, populations, and meta‐populations. We follow this by drawing on examples across a breadth of taxa, and demonstrate that these buffering mechanisms may be quite general. We conclude by identifying key gaps in our knowledge of the fitness and demographic consequences of buffering, in the context of phenological mismatch. Buffering has the potential to substantially alter our understanding of the biotic impacts of future climate change—a greater recognition of the contribution of these mechanisms may reveal that many trophic interactions are surprisingly resilient, and also serve to shift research emphasis to those systems with fewer buffers and towards identifying the limits of those buffers. [ABSTRACT FROM AUTHOR]

Colonization, speciation and extinction are dynamic processes that influence global patterns of species richness.sup.1-6. Island biogeography theory predicts that the contribution of these processes to the accumulation of species diversity depends on the area and isolation of the island.sup.7,8. Notably, there has been no robust global test of this prediction for islands where speciation cannot be ignored.sup.9, because neither the appropriate data nor the analytical tools have been available. Here we address both deficiencies to reveal, for island birds, the empirical shape of the general relationships that determine how colonization, extinction and speciation rates co-vary with the area and isolation of islands. We compiled a global molecular phylogenetic dataset of birds on islands, based on the terrestrial avifaunas of 41 oceanic archipelagos worldwide (including 596 avian taxa), and applied a new analysis method to estimate the sensitivity of island-specific rates of colonization, speciation and extinction to island features (area and isolation). Our model predicts--with high explanatory power--several global relationships. We found a decline in colonization with isolation, a decline in extinction with area and an increase in speciation with area and isolation. Combining the theoretical foundations of island biogeography.sup.7,8 with the temporal information contained in molecular phylogenies.sup.10 proves a powerful approach to reveal the fundamental relationships that govern variation in biodiversity across the planet. Using a global molecular phylogenetic dataset of birds on islands, the sensitivity of island-specific rates of colonization, speciation and extinction to island features (area and isolation) is estimated., Author(s): Luis Valente [sup.1] [sup.2] [sup.3] [sup.4] , Albert B. Phillimore [sup.5] , Martim Melo [sup.6] [sup.7] [sup.8] , Ben H. Warren [sup.9] , Sonya M. Clegg [sup.10] [sup.11] , [...]

The deciduous tree‐herbivorous caterpillar‐insectivorous bird food chain is a well‐studied system for investigating the impacts of climate change across trophic levels. To date, across Europe, most attention has focused on the impacts of increasing spring temperature on changes to phenology in Oak‐dominated (Quercus spp.) woodlands. Paridae species and Pied Flycatcher Ficedula hypoleuca are the most studied secondary consumers, all of which demonstrate an advancement in reproductive phenology with increases in spring temperature. Shifts in climate and phenology may also impact on reproductive investment in clutch size, and the effects of climate on phenology and clutch size may vary depending on woodland composition. To date, the effects of among‐habitat variation in phenology and reproductive investment have received little attention. Insectivorous birds inhabiting woodlands that differ in tree composition may differ in the timing of breeding, due to local tree leafing phenology acting as a cue for egg‐laying date and/or clutch size. Moreover, for most insectivorous birds, woodland composition within a territory is likely to be the main determinant of food availability for both adults and chicks. Consequently, if warming springs affect the temporal patterns of food availability differently across different woodland compositions, this may affect the optimal average local phenology for nesting birds. Here, using data from 34 long‐term (mean 15 years) nest monitoring sites across the UK, we investigate the effect of woodland tree composition and temperature on Blue Tit Cyanistes caeruleus first egg date (FED) and clutch size. We supplemented the nest monitoring data by quantifying woodland composition, at a site level, through modified point counts. We predict that birds breeding in woodlands with greater proportions of late‐leafing species, such as Oak and Ash Fraxinus excelsior, will breed later than those breeding in woodlands with greater proportions of early‐leafing species, such as Birch Betula spp. and Beech Fagus sylvatica. We found no evidence for differences in Blue Tit FED or clutch size in relation to the proportion of any of the tree species investigated, after controlling for temperature and latitude (FED: −3.4 and 2.2, clutch size: −0.4 and − 0.2 eggs for one‐unit increase in temperature and latitude, respectively). In recent decades and across all sites, clutch size has decreased as spring temperatures have increased, a strategy which could allow birds flexibly to adjust their breeding phenology such that nestling demand coincides with peak food availability. The lack of an effect of woodland composition on Blue Tit phenology suggests Blue Tits do not fine‐tune their reproductive phenology to the local tree composition. Whether this lack of evidence for phenological divergence is due to an absence of divergent selection on breeding phenology and clutch size or to gene flow is not clear. [ABSTRACT FROM AUTHOR]

Flowers blooming, fungi fruiting, insects biting, fish spawning, geese migrating, deer calving; our consciousness is steeped in a seasonal calendar of nature's events. Phenology is the study of these recurring, seasonal life-history events, though nowadays this term is widely applied to the events themselves. From Shakespeare's sonnet 98, " From you I have been absent in the spring ", to the appearance of seasonal events and migratory species in the oral traditions of Native Americans, interest in phenology is long-standing and transcends cultures. In this primer we introduce the study of phenology, trace the development of the field, and examine the prominent role phenology has played in evidencing the widespread impacts of anthropogenic climate change on life on Earth. We will consider the potential implications of climatic change for the ability of populations to persist and the stability of species interactions. Phenology, the timing of an organism's recurrent life history events, determines the abiotic conditions and biotic interactions with which it is presented. In this primer, Macphie and Phillimore provide an overview of the science of phenology and discuss the implications of these important timing mechanisms for the ability of populations to persist in the face of climate change. [ABSTRACT FROM AUTHOR]

Advances in spring phenology are among the clearest biological responses to climate warming. There has been much interest in how climate impacts on phenology because the timings of key events have implications for species interactions, nutrient cycling and ecosystem services. To date most work has focused on only one aspect of population phenology, the effects of temperature on the average timing. In comparison, effects of temperature on the abundance of individuals and their seasonal spread are understudied, despite their potential to have profound impacts on species interactions.Here we develop a new method that directly estimates the effect of spring temperatures on the timing, height and width of the phenological distribution and apply it to temperate forest caterpillars, a guild that has been the focus of much research on phenology and trophic mismatch.We find that warmer spring conditions advance the timing of the phenological distribution of abundance by −4.96 days °C−1 and increase its height by 34% °C−1 but have no significant effect on the duration of the distribution. An increase in the maximum density of arboreal caterpillars with rising temperatures has implications for understanding climate impacts on forest food chains, both in terms of herbivory pressure and the resources available to secondary consumers.The new method we have developed allows the thermal sensitivity in the full phenological distribution to be modelled directly from raw data, providing a flexible approach that has broad applicability within global change research. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

In an epoch of rapid environmental change, understanding and predicting how biodiversity will respond to a changing climate is an urgent challenge. Since we seldom have sufficient long‐term biological data to use the past to anticipate the future, spatial climate–biotic relationships are often used as a proxy for predicting biotic responses to climate change over time. These 'space‐for‐time substitutions' (SFTS) have become near ubiquitous in global change biology, but with different subfields largely developing methods in isolation. We review how climate‐focussed SFTS are used in four subfields of ecology and evolution, each focussed on a different type of biotic variable – population phenotypes, population genotypes, species' distributions, and ecological communities. We then examine the similarities and differences between subfields in terms of methods, limitations and opportunities. While SFTS are used for a wide range of applications, two main approaches are applied across the four subfields: spatial in situ gradient methods and transplant experiments. We find that SFTS methods share common limitations relating to (i) the causality of identified spatial climate–biotic relationships and (ii) the transferability of these relationships, i.e. whether climate–biotic relationships observed over space are equivalent to those occurring over time. Moreover, despite widespread application of SFTS in climate change research, key assumptions remain largely untested. We highlight opportunities to enhance the robustness of SFTS by addressing key assumptions and limitations, with a particular emphasis on where approaches could be shared between the four subfields. [ABSTRACT FROM AUTHOR]

Biogeographical factors that influence phenotypic divergence patterns in birds were studied. No significant relationship was found between species age and subspecies richness in birds, however high subspecies richness was found to be associated with large breeding range size.

Birds build nests primarily as a receptacle to lay their eggs in, but they can also provide secondary benefits including structural support, camouflage and adjustment of the microclimate surrounding the eggs and offspring. The factors underlying intraspecific variation in nest characteristics are poorly understood. In this study, we aim to identify the environmental factors that predict nest height variation and the duration of nest building in blue tits Cyanistes caeruleus, evaluating latitude, elevation, temperature and the timing of egg‐laying as predictors of nest height, while also taking into account female and male parental identity. Using 713 nest height observations collected over a period of five years along a 220 km transect in Scotland, we found that if the annual mean timing of egg‐laying was earlier, nests were taller. However, there was no correlation between nest height and elevation, latitude, the minimum temperature in the 14 days pre‐egg‐laying or the phenology of birds within a year. Female parental identity accounted for a large amount of variation in nest height, suggesting that individual behaviour has an influence on nest structure. We also found that nest building duration was shorter when egg laying occurred earlier in the year, and that across all observations taller nests took longer to build. Overall, our results show that blue tits are able to alter their nest characteristics based on environmental gradients like latitude (in the case of building duration) and the annual mean phenological variation of egg laying, and that birds build relatively taller nests faster. [ABSTRACT FROM AUTHOR]

As temperatures rise, timing of reproduction is changing at different rates across trophic levels, potentially resulting in asynchrony between consumers and their resources. The match–mismatch hypothesis (MMH) suggests that trophic asynchrony will have negative impacts on average productivity of consumers. It is also thought to lead to selection on timing of breeding, as the most asynchronous individuals will show the greatest reductions in fitness.Using a 30‐year individual‐level dataset of breeding phenology and success from a population of European shags on the Isle of May, Scotland, we tested a series of predictions consistent with the hypothesis that fitness impacts of trophic asynchrony are increasing.These predictions quantified changes in average annual breeding success and strength of selection on timing of breeding, over time and in relation to rising sea surface temperature (SST) and diet composition.Annual average (population) breeding success was negatively correlated with average lay date yet showed no trend over time, or in relation to increasing SST or the proportion of principal prey in the diet, as would be expected if trophic mismatch was increasing. At the individual level, we found evidence for stabilising selection and directional selection for earlier breeding, although the earliest birds were not the most productive. However, selection for earlier laying did not strengthen over time, or in relation to SST or slope of the seasonal shift in diet from principal to secondary prey. We found that the optimum lay date advanced by almost 4 weeks during the study, and that the population mean lay date tracked this shift.Our results indicate that average performance correlates with absolute timing of breeding of the population, and there is selection for earlier laying at the individual level. However, we found no fitness signatures of a change in the impact of climate‐induced trophic mismatch, and evidence that shags are tracking long‐term shifts in optimum timing. This suggests that if asynchrony is present in this system, breeding success is not impacted. Our approach highlights the advantages of examining variation at both population and individual levels when assessing evidence for fitness impacts of trophic asynchrony. [ABSTRACT FROM AUTHOR]

An accumulating number of studies are reporting severe insect declines. These studies aim to quantify temporal changes in invertebrate populations and community composition and attribute them to anthropogenic drivers.Seibold et al. 2019 (Nature, 574, 671–674) analysed arthropod biomass, abundance and species richness from forest and grassland plots in a region of Germany and reported declines of up to 78% between 2008 and 2018. However, their analysis did not account for the confounding effects of temporal pseudoreplication.We show that simply by including a year random effect in the statistical models and thereby accounting for the common conditions experienced by proximal sites in the same years, four of the five reported declines become non‐significant out of six tests overall.To place recent estimates of insect trends in a broader context, we analysed invertebrate biomass, abundance and richness from 640 time series from 1167 sites around the world. We found that the average trends across the terrestrial and freshwater realms were not significantly distinguishable from no net change. Shorter time series that are likely most affected by sampling error variance – such as those in Seibold et al. 2019 (Nature, 574, 671–674) – yielded the most extreme decline and increase estimates.We suggest that the media uptake of negative trends from short time series may be serving to exaggerate the 'insect Armageddon' and could undermine public confidence in research. We advocate that future research uses appropriate model structures to build a more robust understanding of biodiversity change. [ABSTRACT FROM AUTHOR]

Animal ecologists often collect hierarchically structured data and analyse these with linear mixed‐effects models. Specific complications arise when the effect sizes of covariates vary on multiple levels (e.g. within vs. among subjects). Mean centring of covariates within subjects offers a useful approach in such situations, but is not without problems.A statistical model represents a hypothesis about the underlying biological process. Mean centring within clusters assumes that the lower level responses (e.g. within subjects) depend on the deviation from the subject mean (relative) rather than on the absolute scale of the covariate. This may or may not be biologically realistic. We show that mismatch between the nature of the generating (i.e. biological) process and the form of the statistical analysis produce major conceptual and operational challenges for empiricists.We explored the consequences of mismatches by simulating data with three response‐generating processes differing in the source of correlation between a covariate and the response. These data were then analysed by three different analysis equations. We asked how robustly different analysis equations estimate key parameters of interest and under which circumstances biases arise.Mismatches between generating and analytical equations created several intractable problems for estimating key parameters. The most widely misestimated parameter was the among‐subject variance in response. We found that no single analysis equation was robust in estimating all parameters generated by all equations. Importantly, even when response‐generating and analysis equations matched mathematically, bias in some parameters arose when sampling across the range of the covariate was limited.Our results have general implications for how we collect and analyse data. They also remind us more generally that conclusions from statistical analysis of data are conditional on a hypothesis, sometimes implicit, for the process(es) that generated the attributes we measure. We discuss strategies for real data analysis in face of uncertainty about the underlying biological process. [ABSTRACT FROM AUTHOR]

Asexual reproduction diversifies life‐history priorities and is associated with unusual reproduction and somatic maintenance patterns, such as constant fertility with age, extensive regeneration ability and negligible senescence. While age‐dependent plasticity in relative allocation to sexual versus asexual reproductive modes is relatively well studied, the modulation of somatic maintenance traits in parallel with age‐dependent reproduction is much less well understood in clonal or partially clonal animals.Here, we asked how age‐dependent investment into sexual and asexual reproduction co‐varies with somatic maintenance such as regeneration in a partially clonal freshwater cnidarian Hydra oligactis, a species with remarkable regeneration abilities and experimentally inducible sex.We induced gametogenesis by lowering temperature at two ages, 1 or 4 weeks after detachment from an asexual parent, in animals of a male and a female clone. Then we measured phenotypically asexual and sexual reproductive traits (budding rate, start day and number of sexual organs) together with head regeneration rate, survival and the cellular background of these traits (number of reproductive and interstitial stem cells) for 2 or 5 months.Younger animals had higher asexual reproduction while individuals in the older group had more intensive gametogenesis and reproductive cell production. In parallel with these age‐dependent reproductive differences, somatic maintenance of older individuals was also impacted: head regeneration, survival and interstitial stem cell numbers were reduced compared to younger polyps. Some of the traits investigated showed an ontogenetic effect, suggesting that age‐dependent plasticity and a fixed ontogenetic response might both contribute to differences between age groups.We show that in H. oligactis asexual reproduction coupled with higher somatic maintenance is prioritized earlier in life, while sexual reproduction with higher maintenance costs occurs later if sex is induced. These findings confirm general life‐history theory predictions on resource allocation between somatic maintenance and sexual reproduction applying in a partially clonal species. At the same time, our study also highlights the age‐dependent integration of these resource allocation decisions with sexual/asexual strategies. Accounting for age‐related differences might enhance repeatability of research done with clonal individuals derived from mass cultures. [ABSTRACT FROM AUTHOR]

Hopkins' Bioclimatic Law predicts geographical patterns in phenological timing by establishing a correspondence between latitudinal and altitudinal gradients. First proposed for key phenological events of plants, such as leaf sprouting or flowering dates, this law has rarely been used to assess the geographical equivalence of key life‐history traits of mammals.We hypothesize that (H1) parturition dates of European roe deer Capreolus capreolus are delayed and more synchronized at higher latitudes and altitudes, (H2) parturition timing varies along latitudinal and altitudinal gradients in a way that matches the Hopkins' Bioclimatic Law and (H3) females adjust parturition timing to match the period of high energy demand with peak resource availability.We used parturition dates of 7,444 European roe deer from Switzerland to assess altitudinal variation in birth timing and synchrony from 288 to 2,366 m a.s.l. We then performed a literature survey to compare altitudinal results with those from different populations along the species' latitudinal range of distribution. Finally, we performed spatial analysis combining our highly resolved altitudinal data on parturition dates with plant phenology data.As expected, parturition dates were delayed with increasing latitude and altitude. This delay matched the Bioclimatic Law, as the effect of 1º increase in latitude was similar to 120 m increase in altitude. However, while parturitions were more synchronized with increasing altitude, we did not detect any trend along the latitudinal gradient. Finally, plant phenology explained altitudinal variation in parturition timing better than a linear effect of altitude.Our findings clearly demonstrate the ability of a large herbivore to match parturition timing with phenological conditions across the altitudinal gradient, even at the smallest spatial scales. [ABSTRACT FROM AUTHOR]

Tree diversity exerts a strong influence on consumer communities, but most work has involved single time point measurements over short time periods. Describing temporal variation associated with diversity effects over longer time periods is necessary to fully understand the effects of tree diversity on ecological function.We conducted a year‐long study in an experimental system in southern Mexico assessing the effects of tree diversity on the abundance and diversity of foraging birds. To this end, we recorded bird visitation patterns in 32 tree plots (21 × 21 m; 12 tree species monocultures, 20 four‐species polycultures) every 45 days (n = 8 surveys) and for each plot estimated bird abundance, richness, functional diversity (FD) and phylogenetic diversity (PD). In each case, we reported temporal (intra‐annual) variation in the magnitude of tree diversity effects, and calculated the temporal stability of these bird responses.Across surveys, tree diversity noticeably affected bird responses, demonstrated by significantly higher abundance (43%), richness (32%), PD (25%) and FD (25%) of birds visiting polyculture plots compared to monoculture plots, as well as a distinct species composition between plot types. We also found intra‐annual variation in tree diversity effects on these response variables, ranging from surveys for which the diversity effect was not significant to surveys where a significant 80% increase (e.g. for bird FD and PD) was observed in polyculture relative to monoculture plots. Notably, tree diversity increased the stability of all bird responses, with polycultures having a greater stability abundance (18%), richness (38%), PD (32%), and FD (35%) of birds visiting tree species polycultures compared to monocultures.These results show that tree diversity not only increases bird visitation to plots, but also stabilizes bird habitat usage over time in ways that could implicate insurance‐related mechanisms. Such findings are highly relevant for understanding the long‐term effects of plant diversity on vertebrates and the persistence of bird‐related ecosystem functions. More work is needed to unveil the ecological mechanisms behind temporal variation in vertebrate responses to tree diversity and their consequences for community structure and function. [ABSTRACT FROM AUTHOR]

Rather little is known about the dietary richness and variation of generalist insectivorous species, including birds, due primarily to difficulties in prey identification. Using faecal metabarcoding, we provide the most comprehensive analysis of a passerine's diet to date, identifying the relative magnitudes of biogeographic, habitat and temporal trends in the richness and turnover in diet of Cyanistes caeruleus (blue tit) along a 39 site and 2° latitudinal transect in Scotland. Faecal samples were collected in 2014–2015 from adult birds roosting in nestboxes prior to nest building. DNA was extracted from 793 samples and we amplified COI and 16S minibarcodes. We identified 432 molecular operational taxonomic units that correspond to putative dietary items. Most dietary items were rare, with Lepidoptera being the most abundant and taxon‐rich prey order. Here, we present a statistical approach for estimation of gradients and intersample variation in taxonomic richness and turnover using a generalised linear mixed model. We discuss the merits of this approach over existing tools and present methods for model‐based estimation of repeatability, taxon richness and Jaccard indices. We found that dietary richness increases significantly as spring advances, but changes little with elevation, latitude or local tree composition. In comparison, dietary composition exhibits significant turnover along temporal and spatial gradients and among sites. Our study shows the promise of faecal metabarcoding for inferring the macroecology of food webs, but we also highlight the challenge posed by contamination and make recommendations of laboratory and statistical practices to minimise its impact on inference. [ABSTRACT FROM AUTHOR]

Global climate change has sparked a vast research effort into the demographic and evolutionary consequences of mismatches between consumer and resource phenology. Many studies have used the difference in peak dates to quantify phenological synchrony (match in dates, MD), but this approach has been suggested to be inconclusive, since it does not incorporate the temporal overlap between the phenological distributions (match in overlap, MO).We used 24 years of detailed data on the phenology of a predator–prey system, the great tit (Parus major) and the main food for its nestlings, caterpillars, to estimate MD and MO at the population and brood levels. We compared the performance of both metrics on two key demographic parameters: offspring recruitment probability and selection on the timing of reproduction.Although MD and MO correlated quadratically as expected, MD was a better predictor for both offspring recruitment and selection on timing than MO. We argue—and verify through simulations—that this is because quantifying MO has to be based on nontrivial, difficult‐to‐verify assumptions that likely render MO too inaccurate as a proxy for food availability in practice.Our results have important implications for the allocation of research efforts in long‐term population studies in highly seasonal environments. [ABSTRACT FROM AUTHOR]

A classic system for studying trophic mismatch focuses on the timing of the spring caterpillar peak in relation to the breeding time and productivity of woodland passerine birds. Most work has been conducted in single-site oak woodlands, and little is known about how insights generalize to other woodland types or across space. Here we present the results of a 3-year study on the species composition and temporal distribution of the spring caterpillar peak on different tree taxa across 40 woodland sites spanning 2° of latitude in Scotland. We used molecular barcoding to identify 62 caterpillar species, with winter moth (Operophtera brumata) being the most abundant, comprising one-third of the sample. Oak (Quercus sp.) and willow (Salix sp.) hosted significantly higher caterpillar abundances than other tree taxa, with winter moth exhibiting similar trends and invariantly proportionate across tree taxa. Caterpillar peak phenology was broadly similar between tree taxa. While latitude had little effect, increasing elevation increased the height of the caterpillar peak and retarded timing by 3.7 days per 100 m. These findings extend our understanding of how mismatch may play out spatially, with caterpillar peak date varying with elevation and tree taxa varying in the caterpillar resource that they host. [ABSTRACT FROM AUTHOR]

A phylogenetic comparative approach is used for testing the affect of novel environments versus insularity on rates of body size and sexual size dimorphism diversification in Anolis. The elevated diversifying selection due to ecological release has driven high rates of body size diversification in both small-island and mainland novel environments.

Establishing the cues or constraints that influence avian timing of breeding is the key to accurate prediction of future phenology. This study aims to identify the aspects of the environment that predict the timing of two measures of breeding phenology (nest initiation and egg laying date) in an insectivorous woodland passerine, the blue tit (Cyanistes caeruleus). We analyse data collected from a 220 km, 40-site transect over 3 years and consider spring temperatures, tree leafing phenology, invertebrate availability and photoperiod as predictors of breeding phenology. We find that mean night-time temperature in early spring is the strongest predictor of both nest initiation and lay date and suggest this finding is most consistent with temperature acting as a constraint on breeding activity. Birch budburst phenology significantly predicts lay date additionally to temperature, either as a direct cue or indirectly via a correlated variable. We use cross-validation to show that our model accurately predicts lay date in two further years and find that similar variables predict lay date well across the UK national nest record scheme. This work refines our understanding of the principal factors influencing the timing of tit reproductive phenology and suggests that temperature may have both a direct and indirect effect. [ABSTRACT FROM AUTHOR]

The Arctic is undergoing dramatic environmental change with rapidly rising surface temperatures, accelerating sea ice decline and changing snow regimes, all of which influence tundra plant phenology. Despite these changes, no globally consistent direction of trends in spring phenology has been reported across the Arctic. While spring has advanced at some sites, spring has delayed or not changed at other sites, highlighting substantial unexplained variation. Here, we test the relative importance of local temperatures, local snow melt date and regional spring drop in sea ice extent as controls of variation in spring phenology across different sites and species. Trends in long‐term time series of spring leaf‐out and flowering (average span: 18 years) were highly variable for the 14 tundra species monitored at our four study sites on the Arctic coasts of Alaska, Canada and Greenland, ranging from advances of 10.06 days per decade to delays of 1.67 days per decade. Spring temperatures and the day of spring drop in sea ice extent advanced at all sites (average 1°C per decade and 21 days per decade, respectively), but only those sites with advances in snow melt (average 5 days advance per decade) also had advancing phenology. Variation in spring plant phenology was best explained by snow melt date (mean effect: 0.45 days advance in phenology per day advance snow melt) and, to a lesser extent, by mean spring temperature (mean effect: 2.39 days advance in phenology per °C). In contrast to previous studies examining sea ice and phenology at different spatial scales, regional spring drop in sea ice extent did not predict spring phenology for any species or site in our analysis. Our findings highlight that tundra vegetation responses to global change are more complex than a direct response to warming and emphasize the importance of snow melt as a local driver of tundra spring phenology. [ABSTRACT FROM AUTHOR]

Widespread population declines have been reported for diverse Mediterranean butterflies over the last three decades, and have been significantly associated with increased global change impacts. The specific landscape and climatic drivers of these declines remain uncertain for most declining species.Here, we analyse whether plastic phenotypic traits of a model butterfly species (Pieris napi) perform as reliable biomarkers of vulnerability to extreme temperature impacts in natural populations, showing contrasting trends in thermally exposed and thermally buffered populations.We also examine whether improved descriptions of thermal exposure of insect populations can be achieved by combining multiple information sources (i.e., integrating measurements of habitat thermal buffering, habitat thermal amplification, host plant transpiration, and experimental assessments of thermal death time (TDT), thermal avoidance behaviour (TAB) and thermally induced trait plasticity). These integrative analyses are conducted in two demographically declining and two non‐declining populations of P. napi.The results show that plastic phenotypic traits (butterfly body mass and wing size) are reliable biomarkers of population vulnerability to extreme thermal conditions. Butterfly wing size is strongly reduced only in thermally exposed populations during summer drought periods. Laboratory rearing of these populations documented reduced wing size due to significant negative effects of increased temperatures affecting larval growth. We conclude that these thermal biomarkers are indicative of the population vulnerability to increasing global warming impacts, showing contrasting trends in thermally exposed and buffered populations.Thermal effects in host plant microsites significantly differ between populations, with stressful thermal conditions only effectively ameliorated in mid‐elevation populations. In lowland populations, we observe a sixfold reduction in vegetation thermal buffering effects, and larval growth occurs in these populations at significantly higher temperatures. Lowland populations show reduced host plant quality (C/N ratio), reduced leaf transpiration rates and complete above‐ground plant senescence during the peak of summer drought. Amplified host plant temperatures are observed in open microsites, reaching thermal thresholds that can affect larval survival.Overall, our results suggest that butterfly population vulnerability to long‐term drought periods is associated with multiple co‐occurring and interrelated ecological factors, including limited vegetation thermal buffering effects at lowland sites, significant drought impacts on host plant transpiration and amplified leaf surface temperature, as well as reduced leaf quality linked to the seasonal advance of plant phenology. Our results also identify multiannual summer droughts affecting larval growing periods as a key driver of the recently reported butterfly population declines in the Mediterranean biome. Plastic phenotypic traits are reliable biomarkers of population vulnerability to extreme thermal conditions. Population vulnerability to long‐term drought periods is associated with multiple co‐occurring factors, including vegetation thermal buffering effects, drought impacts on host plant transpiration and amplified leaf surface temperature, as well as reduced leaf quality linked to plant phenology. [ABSTRACT FROM AUTHOR]

The decline of farmland birds across Europe is a well‐documented case of biodiversity loss, and despite land stewardship supported by funding from agri‐environment schemes (AES), the negative trends have not yet been reversed.To investigate the contribution of AES towards farmland bird conservation, we compared abundance of five farmland bird species across 13 years and 53 farms (158 farm years = AES, 72 farm years = non AES) in Northeastern Scotland (UK), a region with relatively mixed farmland.Between 2003 and 2015, on both AES and control farms, skylark (Alauda arvensis) showed a nonsignificant decline, and tree sparrow (Passer montanus) and yellowhammer (Emberiza citrinella) nonsignificant increases, whereas reed bunting (Emberiza schoeniclus) and linnet (Carduelis cannabina) populations remained relatively stable.We did not detect a significant association between AES and avian abundance or population trends for any of these species, but there were positive associations with some AES management options.Possible explanations for the lack of a significant AES‐bird abundance association include poor uptake of the best AES options for farmland birds, suboptimal implementation, spill‐over effects from AES onto control farms, and the relatively good state of farmland habitats outwith AES in Northeastern Scotland.Synthesis and applications. We documented a weak effect size of participation in agri‐environment schemes on farmland bird abundance. We therefore recommend future monitoring studies be designed after consulting a power analysis. Among different land management options, we found that species‐rich grasslands, water margins, and wetland creation enhanced breeding bird abundance, highlighting the importance of relatively undisturbed herbaceous or grassland vegetation for farmland conservation. We documented a weak effect size of participation in agri‐environment schemes on farmland bird abundance. We therefore recommend future monitoring studies be designed after consulting a power analysis. Among different land management options, we found that species‐rich grasslands, water margins, and wetland creation enhanced breeding bird abundance, highlighting the importance of relatively undisturbed herbaceous or grassland vegetation for farmland conservation. [ABSTRACT FROM AUTHOR]

The nesting phenology and productivity of hole‐nesting woodland passerines, such as tit species (Paridae), has been the subject of many studies and played a central role in advancing our understanding of the causes and consequences of trophic mismatch. However, as most studies have been conducted in mature, oak‐rich (Quercus sp.) woodlands, it is unknown whether insights from such studies generalise to other habitats used by woodland generalist species. Here we applied spatial mixed models to data collected over three years (2014–2016) from 238 nestboxes across 40 sites – that vary in woodland habitat and elevation – along a 220 km transect in Scotland. We evaluate the importance of habitat, biogeography and food availability as predictors of mesoscale among‐site variation in blue tit Cyanistes caeruleus nestbox occupancy and two components of productivity (clutch size and fledging success). We found that habitat was not a significant predictor of occupancy or clutch size but that occupancy exhibited pronounced biogeographic trends, declining with increasing latitude and elevation. However, fledging success, defined as the proportion of a clutch that fledged, was positively correlated with site level availability of birch, oak and sycamore, and tree diversity. The lack of correspondence between the effects of habitat on fledging success versus occupancy and clutch size may indicate that blue tits do not accurately predict the future quality of their breeding sites when selecting territories and laying clutches. We found little evidence of spatial autocorrelation in occupancy or clutch size, whereas spatial autocorrelation in fledging success extends over multiple sites, albeit non‐significantly. Taken together, our findings suggest that the relationship between breeding decisions and breeding outcomes varies among habitats, and we urge caution when extrapolating inferences from one habitat to others. [ABSTRACT FROM AUTHOR]

One consequence of rising spring temperatures is that the optimum timing of key life-history events may advance. Where this is the case, a population's fate may depend on the degree to which it is able to track a change in the optimum timing either via plasticity or via adaptation. Estimating the effect that temperature change will have on optimum timing using standard approaches is logistically challenging, with the result that very few estimates of this important parameter exist. Here we adopt an alternative statistical method that substitutes space for time to estimate the temperature sensitivity of the optimum timing of 22 plant species based on >200 000 spatiotemporal phenological observations from across the United Kingdom. We find that first leafing and flowering dates are sensitive to forcing (spring) temperatures, with optimum timing advancing by an average of 3 days °C−1 and plastic responses to forcing between −3 and −8 days °C−1. Chilling (autumn/winter) temperatures and photoperiod tend to be important cues for species with early and late phenology, respectively. For most species, we find that plasticity is adaptive, and for seven species, plasticity is sufficient to track geographic variation in the optimum phenology. For four species, we find that plasticity is significantly steeper than the optimum slope that we estimate between forcing temperature and phenology, and we examine possible explanations for this countergradient pattern, including local adaptation. [ABSTRACT FROM AUTHOR]

Understanding and predicting phenology has become more important with ongoing climate change and has brought about great research efforts in the recent decades. The majority of studies examining spring phenology of insects have focussed on the effects of spring temperatures alone., Here we use citizen-collected observation data to show that winter cold duration, in addition to spring temperature, can affect the spring emergence of butterflies. Using spatial mixed models, we disentangle the effects of climate variables and reveal impacts of both spring and winter conditions for five butterfly species that overwinter as pupae across the UK, with data from 1976 to 2013 and one butterfly species in Sweden, with data from 2001 to 2013., Warmer springs lead to earlier emergence in all species and milder winters lead to statistically significant delays in three of the five investigated species. We also find that the delaying effect of winter warmth has become more pronounced in the last decade, during which time winter durations have become shorter., For one of the studied species, Anthocharis cardamines (orange tip butterfly), we also make use of parameters determined from previous experiments on pupal development to model the spring phenology. Using daily temperatures in the UK and Sweden, we show that recent variation in spring temperature corresponds to 10-15 day changes in emergence time over UK and Sweden, whereas variation in winter duration corresponds to 20 days variation in the south of the UK versus only 3 days in the south of Sweden., In summary, we show that short winters delay phenology. The effect is most prominent in areas with particularly mild winters, emphasising the importance of winter for the response of ectothermic animals to climate change. With climate change, these effects may become even stronger and apply also at higher latitudes. [ABSTRACT FROM AUTHOR]

Predicting habitat quality is a major challenge for animals selecting a breeding patch, because it affects reproductive success. Breeding site selection may be based on previous experience, or on social information from the density and success of competitors with an earlier phenology., Variation in animal breeding phenology is often correlated with variation in habitat quality. Generally, animals breed earlier in high-quality habitats that allow them to reach a nutritional threshold required for breeding earlier or avoid nest predation. In addition, habitat quality may affect phenological overlap between species and thereby interspecific competition. Therefore, we hypothesized that competitor breeding phenology can be used as social cue by settling migrants to locate high-quality breeding sites., To test this hypothesis, we experimentally advanced and delayed hatching phenology of two resident tit species on the level of study plots and studied male and female settlement patterns of migratory pied flycatchers Ficedula hypoleuca. The manipulations were assigned at random in two consecutive years, and treatments were swapped between years in sites that were used in both years., In both years, males settled in equal numbers across treatments, but later arriving females avoided pairing with males in delayed phenology plots. Moreover, male pairing probability declined strongly with arrival date on the breeding grounds., Our results demonstrate that competitor phenology may be used to assess habitat quality by settling migrants, but we cannot pinpoint the exact mechanism (e.g. resource quality, predation pressure or competition) that has given rise to this pattern., In addition, we show that opposing selection pressures for arrival timing may give rise to different social information availabilities between sexes. We discuss our findings in the context of climate warming, social information use and the evolution of protandry in migratory animals. [ABSTRACT FROM AUTHOR]

Projecting the fates of populations under climate change is one of global change biology's foremost challenges. Here, we seek to identify the contributions that temperature-mediated local adaptation and plasticity make to spatial variation in nesting phenology, a phenotypic trait showing strong responses to warming. We apply a mixed modeling framework to a Britain-wide spatiotemporal dataset comprising >100 000 records of first egg dates from four single-brooded passerine bird species. The average temperature during a specific time period (sliding window) strongly predicts spatiotemporal variation in lay date. All four species exhibit phenological plasticity, advancing lay date by 2-5 days °C−1. The initiation of this sliding window is delayed further north, which may be a response to a photoperiod threshold. Using clinal trends in phenology and temperature, we are able to estimate the temperature sensitivity of selection on lay date ( B), but our estimates are highly sensitive to the temporal position of the sliding window. If the sliding window is of fixed duration with a start date determined by photoperiod, we find B is tracked by phenotypic plasticity. If, instead, we allow the start and duration of the sliding window to change with latitude, we find plasticity does not track B, although in this case, at odds with theoretical expectations, our estimates of B differ across latitude vs. longitude. We argue that a model combining photoperiod and mean temperature is most consistent with current understanding of phenological cues in passerines, the results from which suggest that each species could respond to projected increases in spring temperatures through plasticity alone. However, our estimates of B require further validation. [ABSTRACT FROM AUTHOR]