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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]

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]

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]

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]

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]

Although the number of studies discerning the impact of climate change on ecological systems continues to increase, there has been relatively little sharing of the lessons learnt when accumulating this evidence. At a recent workshop entitled 'Using climate data in ecological research' held at the UK Met Office, ecologists and climate scientists came together to discuss the robust analysis of climate data in ecology. The discussions identified three common pitfalls encountered by ecologists: 1) selection of inappropriate spatial resolutions for analysis; 2) improper use of publically available data or code; and 3) insufficient representation of the uncertainties behind the adopted approach. Here, we discuss how these pitfalls can be avoided, before suggesting ways that both ecology and climate science can move forward. Our main recommendation is that ecologists and climate scientists collaborate more closely, on grant proposals and scientific publications, and informally through online media and workshops. More sharing of data and code (e.g. via online repositories), lessons and guidance would help to reconcile differing approaches to the robust handling of data. We call on ecologists to think critically about which aspects of the climate are relevant to their study system, and to acknowledge and actively explore uncertainty in all types of climate data. And we call on climate scientists to make simple estimates of uncertainty available to the wider research community. Through steps such as these, we will improve our ability to robustly attribute observed ecological changes to climate or other factors, while providing the sort of influential, comprehensive analyses that efforts to mitigate and adapt to climate change so urgently require. [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]

There are wide reports of advances in the timing of spring migration of birds over time and in relation to rising temperatures, though phenological responses vary substantially within and among species. An understanding of the ecological, life-history and geographic variables that predict this intra- and interspecific variation can guide our projections of how populations and species are likely to respond to future climate change., Here, we conduct phylogenetic meta-analyses addressing slope estimates of the timing of avian spring migration regressed on (i) year and (ii) temperature, representing a total of 413 species across five continents. We take into account slope estimation error and examine phylogenetic, ecological and geographic predictors of intra- and interspecific variation., We confirm earlier findings that on average birds have significantly advanced their spring migration time by 2·1 days per decade and 1·2 days °C−1. We find that over time and in response to warmer spring conditions, short-distance migrants have advanced spring migratory phenology by more than long-distance migrants. We also find that larger bodied species show greater advance over time compared to smaller bodied species. Our results did not reveal any evidence that interspecific variation in migration response is predictable on the basis of species' habitat or diet., We detected a substantial phylogenetic signal in migration time in response to both year and temperature, suggesting that some of the shifts in migratory phenological response to climate are predictable on the basis of phylogeny. However, we estimate high levels of species and spatial variance relative to phylogenetic variance, which is consistent with plasticity in response to climate evolving fairly rapidly and being more influenced by adaptation to current local climate than by common descent., On average, avian spring migration times have advanced over time and as spring has become warmer. While we are able to identify predictors that explain some of the true among-species variation in response, substantial intra- and interspecific variation in migratory response remains to be explained. [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]

Slowdowns in lineage accumulation in phylogenies suggest that speciation rates decline as diversity increases. Likelihood methods have been developed to detect such diversity dependence. However, a thorough test of whether such approaches correctly infer diversity dependence is lacking., Here, we simulate phylogenetic branching under linear negative diversity-dependent and diversity-independent models and estimate from the simulated phylogenies the maximum-likelihood parameters for three different conditionings - on survival of the birth-death process given the crown age, on tree size ( N) and on tree size given the crown age. We report the accuracy of recovering the simulation parameters and the reliability of the model selection based on the χ2 likelihood ratio test., Parameter estimate accuracy: Conditioning on survival given the crown age yields a severe bias of the carrying capacity K towards N and an upward bias of the speciation rate, particularly in clades where diversity-dependent feedbacks are still weak ( N « K). Conditioning on N yields an overestimate of K and an underestimate of speciation rate, particularly when saturation has been reached. Dual conditioning yields relatively unbiased parameter estimates on average, but the deviation from the true value for any single estimate may be large., Model selection reliability: The frequency of incorrectly rejecting a diversity-independent model when the simulation was diversity-independent (type I error) differs substantially from the significance level α used in the likelihood ratio test, rendering the likelihood ratio test inappropriate. The frequency of correctly rejecting the diversity-independent model when the simulation was diversity-dependent (power) is larger when the clade is closer to equilibrium and for conditioning on crown age., We conclude that conditioning on crown age has the best statistical properties overall, but caution that parameter estimates may be biased. To assess parameter uncertainty in future studies of diversity dependence on real data, we recommend parametric bootstrapping, examination of the likelihood surface and comparison of estimates across the types of conditioning. To assess model selection reliability, we discourage the use of the χ2 likelihood ratio test or AIC (which are equivalent in this case), but recommend a likelihood ratio test based on parametric bootstrap. We illustrate this method for the diversification of Dendroica warblers. [ABSTRACT FROM AUTHOR]

Climate change may cause phenological asynchrony between trophic levels, which can lead to mismatched reproduction in animals. Although indirect effects of mismatch on fitness are well described, direct effects on parental prey choice are not. Moreover, direct effects of prey variation on offspring condition throughout their early development are understudied., Here, we used camera trap data collected over 2 years to study the effects of trophic mismatch and nestling age on prey choice in pied flycatchers ( Ficedula hypoleuca). Furthermore, we studied the effect of mismatch and variation in nestling diet on offspring condition., Both experimentally induced and natural mismatches with the caterpillar peak negatively affected absolute and relative numbers of caterpillars and offspring condition (mass, tarsus and wing length) and positively affected absolute and relative numbers of flying insects in the nestling diet. Feeding more flying insects was negatively correlated with nestling day 12 mass., Both descriptive and experimental data showed preferential feeding of spiders when nestlings were <7 days old. Receiving more spiders during this phase was positively correlated with tarsus growth., These results highlight the need for a more inclusive framework to study phenological mismatch in nature. The general focus on only one prey type, the rarity of studies that measure environmental abundance of prey, and the lack of timing experiments in dietary studies currently hamper understanding of the actual trophic interactions that affect fitness under climate change. [ABSTRACT FROM AUTHOR]

As populations shift their ranges in response to global change, local species assemblages can change, setting the stage for new ecological interactions, community equilibria and evolutionary responses., Here, we focus on the range dynamics of four avian brood parasite species and their hosts in southern Africa, in a context of bush encroachment (increase in woody vegetation density in places previously occupied by savanna-grassland mosaics) favouring some species at the expense of others. We first tested whether hosts and parasites constrained each other's ability to expand or maintain their ranges. Secondly, we investigated whether range shifts represented an opportunity for new host-parasite and parasite-parasite interactions., We used multispecies dynamic occupancy models with interactions, fitted to citizen science data, to estimate the contribution of interspecific interactions to range shifts and to quantify the change in species co-occurrence probability over a 25-year period., Parasites were able to track their hosts' range shifts. We detected no deleterious effect of the parasites' presence on either the local population viability of host species or the hosts' ability to colonize newly suitable areas. In the recently diversified indigobird radiation ( Vidua spp.), following bush encroachment, the new assemblages presented more potential opportunities for speciation via host switch, but also more potential for hybridization between extant lineages, also via host switch., Multispecies dynamic occupancy models with interactions brought new insights into the feedbacks between range shifts, biotic interactions and local demography: brood parasitism had little detected impact on extinction or colonization processes, but inversely the latter processes affected biotic interactions via the modification of co-occurrence patterns. [ABSTRACT FROM AUTHOR]

Male-biased sex ratios occur in many bird species, particularly in those with small or declining populations, but the causes of these skews and their consequences for local population demography are rarely known. Within-species variation in sex ratios can help to identify the demographic and behavioural processes associated with such biases., Small populations may be more likely to have skewed sex ratios if sex differences in survival, recruitment or dispersal vary with local abundance. Analyses of species with highly variable local abundances can help to identify these mechanisms and the implications for spatial variation in demography. Many migratory bird species are currently undergoing rapid and severe declines in abundance in parts of their breeding ranges and thus have sufficient spatial variation in abundance to explore the extent of sex ratio biases, their causes and implications., Using national-scale bird ringing data for one such species (willow warbler, Phylloscopus trochilus), we show that sex ratios vary greatly across Britain and that male-biased sites are more frequent in areas of low abundance, which are now widespread across much of south and east England. These sex ratio biases are sufficient to impact local productivity, as the relative number of juveniles caught at survey sites declines significantly with increasing sex ratio skew., Sex differences in survival could influence this sex ratio variation, but we find little evidence for sex differences in survival increasing with sex ratio skew. In addition, sex ratios have become male-biased over the last two decades, but there are no such trends in adult survival rates for males or females. This suggests that lower female recruitment into low abundance sites is contributing to these skews., These findings suggest that male-biased sex ratios in small and declining populations can arise through local-scale sex differences in survival and dispersal, with females recruiting disproportionately into larger populations. Given the high level of spatial variation in population declines and abundance of many migratory bird species across Europe at present, male-biased small populations may be increasingly common. As singing males are the primary records used in surveys of these species, and as unpaired males often sing throughout the breeding season, local sex ratio biases could also be masking the true extent of these population declines. [ABSTRACT FROM AUTHOR]

The adaptability of species' climatic niches can influence the dynamics of colonization and gene flow across climatic gradients, potentially increasing the likelihood of speciation or reducing extinction in the face of environmental change. However, previous comparative studies have tested these ideas using geographically, taxonomically and ecologically restricted samples, yielding mixed results, and thus the processes linking climatic niche evolution with diversification remain poorly understood., Focusing on birds, the largest and most widespread class of terrestrial vertebrates, we test whether variation in species diversification among clades is correlated with rates of climatic niche evolution and the extent to which these patterns are modified by underlying gradients in biogeography and species' ecology., We quantified climatic niches, latitudinal distribution and ecological traits for 7657 (˜75%) bird species based on geographical range polygons and then used Bayesian phylogenetic analyses to test whether niche evolution was related to species richness and rates of diversification across genus- and family-level clades., We found that the rate of climatic niche evolution has a positive linear relationship with both species richness and diversification rate at two different taxonomic levels (genus and family). Furthermore, this positive association between labile climatic niches and diversification was detected regardless of variation in clade latitude or key ecological traits., Our findings suggest either that rapid adaptation to unoccupied areas of climatic niche space promotes avian diversification, or that diversification promotes adaptation. Either way, we propose that climatic niche evolution is a fundamental process regulating the link between climate and biodiversity at global scales, irrespective of the geographical and ecological context of speciation and extinction. [ABSTRACT FROM AUTHOR]

In seasonal climates, dormancy is a common strategy that structures biodiversity and is necessary for the persistence of many species. Climate change will likely alter dormancy dynamics in zooplankton, the basis of aquatic food webs, by altering two important hatching cues: mean temperatures during the ice-free season, and mean day length when lakes become ice free. Theory suggests that these changes could alter diversity, hatchling abundances and phenology within lakes, and that these responses may diverge across latitudes due to differences in optimal hatching cues and strategies., To examine the role of temperature and day length on hatching dynamics, we collected sediment from 25 lakes across a 1800 km latitudinal gradient and exposed sediment samples to a factorial combination of two photoperiods (12 and 16 h) and two temperatures (8 and 12 °C) representative of historical southern (short photoperiod, warm) and northern (long photoperiod, cool) lake conditions. We tested whether sensitivity to these hatching cues varies by latitudinal origin and differs among taxa., Higher temperatures advanced phenology for all taxa, and these advances were greatest for cladocerans followed by copepods and rotifers. Although phenology differed among taxa, the effect of temperature did not vary with latitude. The latitudinal origin of the egg bank influenced egg abundance and hatchling abundance and diversity, with these latter effects varying with taxa, temperature and photoperiod., Copepod hatchling abundances peaked at mid-latitudes in the high temperature and long photoperiod treatments, whereas hatchling abundances of other zooplankton were greatest at low latitudes and high temperature. The overall diversity of crustacean zooplankton (copepods and cladocerans) also reflected distinct responses of each taxa to our treatments, with the greatest diversity occurring at mid-latitudes (~56 °N) in the shorter photoperiod treatment., Our results demonstrate that hatching cues differ for broad taxonomic groups that vary in developmental and life-history strategies. These differences are predicted to drive latitude-specific shifts in zooplankton emergence with climate change and could alter the base of aquatic food webs. [ABSTRACT FROM AUTHOR]

Climate change alters the frequency and severity of extreme events, such as drought. Such events will be increasingly important in shaping communities as climate change intensifies. The ability of species to withstand extreme events (resistance) and to recover once adverse conditions abate (resilience) will determine their persistence., We estimated the resistance and resilience of bird species during and after a 13-year drought (the 'Big Dry') in floodplain forests in south-eastern Australia., We conducted bird surveys at the beginning and end of the Big Dry, and after the abrupt end to the drought (the 'Big Wet'), to evaluate species-specific changes in reporting rates among the three periods. We assessed changes in bird-breeding activity before and after the Big Wet to estimate demographic resilience based on breeding., Between the start and the end of the Big Dry (1998 vs. 2009), 37 of 67 species declined substantially. Of those, only two had increased reporting rates after the Big Wet (2009 vs. 2013) that were equal to or larger than their declines, while three partially recovered. All other declining species showed low resilience: 25 showed no change in reporting rates and seven declined further. The number of breeding species and total breeding activity of all species declined after the Big Wet, and there was no change in the number of young produced., The Big Dry caused widespread declines in the floodplain avifauna. Despite the drought being broken by 2 years of well-above-average rainfall and subsequent near-average rainfall, most species showed low resilience and there was little indication that overall breeding had increased. The effects of drought appeared to be pervasive for much of the floodplain avifauna, regardless of species traits (species body mass, fecundity, mobility or diet). Ecosystems such as these are likely to require active management and restoration, including reinstatement of natural flooding regimes, to improve ecological condition, to enhance resistance and resilience to extreme climate events. [ABSTRACT FROM AUTHOR]

Phenology shifts are the most widely cited examples of the biological impact of climate change, yet there are few assessments of potential effects on the fitness of individual organisms or the persistence of populations. Despite extensive evidence of climate-driven advances in phenological events over recent decades, comparable patterns across species' geographic ranges have seldom been described. Even fewer studies have quantified concurrent spatial gradients and temporal trends between phenology and climate. Here we analyse a large data set (~129 000 phenology measures) over 37 years across the UK to provide the first phylogenetic comparative analysis of the relative roles of plasticity and local adaptation in generating spatial and temporal patterns in butterfly mean flight dates. Although populations of all species exhibit a plastic response to temperature, with adult emergence dates earlier in warmer years by an average of 6.4 days per °C, among-population differences are significantly lower on average, at 4.3 days per °C. Emergence dates of most species are more synchronised over their geographic range than is predicted by their relationship between mean flight date and temperature over time, suggesting local adaptation. Biological traits of species only weakly explained the variation in differences between space-temperature and time-temperature phenological responses, suggesting that multiple mechanisms may operate to maintain local adaptation. As niche models assume constant relationships between occurrence and environmental conditions across a species' entire range, an important implication of the temperature-mediated local adaptation detected here is that populations of insects are much more sensitive to future climate changes than current projections suggest. [ABSTRACT FROM AUTHOR]

Island biotas emerge from the interplay between colonisation, speciation and extinction and are often the scene of spectacular adaptive radiations. A common assumption is that insular diversity is at a dynamic equilibrium, but for remote islands, such as Hawaii or Galápagos, this idea remains untested. Here, we reconstruct the temporal accumulation of terrestrial bird species of the Galápagos using a novel phylogenetic method that estimates rates of biota assembly for an entire community. We show that species richness on the archipelago is in an ascending phase and does not tend towards equilibrium. The majority of the avifauna diversifies at a slow rate, without detectable ecological limits. However, Darwin's finches form an exception: they rapidly reach a carrying capacity and subsequently follow a coalescent-like diversification process. Together, these results suggest that avian diversity of remote islands is rising, and challenge the mutual exclusivity of the non-equilibrium and equilibrium ecological paradigms. [ABSTRACT FROM AUTHOR]

The rise in spring temperatures over the past half-century has led to advances in the phenology of many nontropical plants and animals. As species and populations differ in their phenological responses to temperature, an increase in temperatures has the potential to alter timing-dependent species interactions. One species-interaction that may be affected is the competition for light in deciduous forests, where early vernal species have a narrow window of opportunity for growth before late spring species cast shade. Here we consider the Marsham phenology time series of first leafing dates of thirteen tree species and flowering dates of one ground flora species, which spans two centuries. The exceptional length of this time series permits a rare comparison of the statistical support for parameter-rich regression and mechanistic thermal sensitivity phenology models. While mechanistic models perform best in the majority of cases, both they and the regression models provide remarkably consistent insights into the relative sensitivity of each species to forcing and chilling effects. All species are sensitive to spring forcing, but we also find that vernal and northern European species are responsive to cold temperatures in the previous autumn. Whether this sensitivity reflects a chilling requirement or a delaying of dormancy remains to be tested. We then apply the models to projected future temperature data under a fossil fuel intensive emissions scenario and predict that while some species will advance substantially others will advance by less and may even be delayed due to a rise in autumn and winter temperatures. Considering the projected responses of all fourteen species, we anticipate a change in the order of spring events, which may lead to changes in competitive advantage for light with potential implications for the composition of temperate forests. [ABSTRACT FROM AUTHOR]

A recent publication (Pedreschi et al., 2014, Journal of Biogeography, 41, 548-560) casts doubt over the status of pike (Esox lucius) as a non-native species in Ireland by reporting two distinct genetic groups of pike present: one a human introduction in the Middle Ages, the other hypothesized to result from natural colonization after the Last Glacial Maximum (LGM). While the existence of two groups is not disputed, the hypothesized natural colonization scenario conflicts with the sequence in which the islands of Britain and Ireland became isolated from Europe after the LGM. An alternative natural colonization scenario raised herein was rejected, leaving an earlier, two-phase, human introduction of pike from Britain or Europe to Ireland as a realistic alternative hypothesis explaining the results of Pedreschi et al. (2014). This leaves the debates on human introduction versus natural colonization, introduced versus native species status, and pike management in Ireland wide open. [ABSTRACT FROM AUTHOR]

Aim One prominent explanation for the latitudinal gradient in biodiversity proposes that its prime cause is the greater age and/or higher origination rates of tropical clades, and the infrequent or delayed dispersal of their component species into temperate regions. An alternative is that species' carrying capacities vary regionally, which influences rates of time-averaged diversification via ecological opportunity. We contrast these hypotheses, in order to assess potential historical influences upon the latitudinal gradient of New World passerine birds (order Passeriformes), comparing patterns among the two suborders present (oscines and suboscines), which are known to have had different routes of dispersal across the region. Location New World. Methods We examine diversity patterns, their abiotic and biotic correlates, and the distributions of phylogenetically old and young species. Results Strong latitudinal gradients are present within both oscine and suboscine birds, with maximum diversity towards the equator, but their overall shapes differ. Among the oscines, older lineages are found towards the north, with progressively younger lineages present further south. Regional variation in oscine richness is statistically well explained by a combination of productivity and elevation ( R2 = 0.76). In contrast, few suboscine groups have colonized the north, so their current diversity is well correlated with temperature seasonality ( R2 = 0.74). Main conclusions Because the oscines colonized the Americas from the north, their latitudinal gradient must reflect regional differences in time-averaged diversification rates, and not the time present within a region. The richness patterns derived from phylogenetic data and the strong predictive power of the normalized difference vegetation index (NDVI) suggest that the radiation of the oscines is consistent with the idea that entry into a new region stimulates a burst of diversification, which is higher and/or continues for longer in areas with greater carrying capacity. Conversely, the suboscine distributions potentially reflect a large historical barrier to dispersal and niche conservatism of climatic tolerances, possibly coupled with competition from the oscines. Although contemporary conditions can explain much of the passerine diversity patterns, history has had an important influence on the taxonomic composition of this gradient. [ABSTRACT FROM AUTHOR]

Aim We investigated genetic variation of Irish pike populations and their relationship with European outgroups, in order to elucidate the origin of this species to the island, which is largely assumed to have occurred as a human-mediated introduction over the past few hundred years. We aimed thereby to provide new insights into population structure to improve fisheries and biodiversity management in Irish freshwaters. Location Ireland, Britain and continental Europe. Methods A total of 752 pike ( Esox lucius) were sampled from 15 locations around Ireland, and 9 continental European sites, and genotyped at six polymorphic microsatellite loci. Patterns and mechanisms of population genetic structure were assessed through a diverse array of methods, including Bayesian clustering, hierarchical analysis of molecular variance, and approximate Bayesian computation. Results Varying levels of genetic diversity and a high degree of population genetic differentiation were detected. Clear substructure within Ireland was identified, with two main groups being evident. One of the Irish populations showed high similarity with British populations. The other, more widespread, Irish strain did not group with any European population examined. Approximate Bayesian computation suggested that this widespread Irish strain is older, and may have colonized Ireland independently of humans. Main conclusions Population genetic substructure in Irish pike is high and comparable to the levels observed elsewhere in Europe. A comparison of evolutionary scenarios upholds the possibility that pike may have colonized Ireland in two 'waves', the first of which, being independent of human colonization, would represent the first evidence for natural colonization of a non-anadromous freshwater fish to the island of Ireland. Although further investigations using comprehensive genomic techniques will be necessary to confirm this, the present results warrant a reappraisal of current management strategies for this species. [ABSTRACT FROM AUTHOR]

Aim The challenge for phylogeography, like other observational sciences, is to extract general relationships representing causes and effects from complex natural data. I describe comparisons of synchronously diverging co-distributed ( SDC) taxa, including sympatric sister species, to help meet this challenge. Location Global. Methods As an example, using narrative and best evidence synthesis, I re-evaluate ad hoc aggregate analyses relating population genetic structure ( FST) to dispersal potential. I deconstruct an aggregate global analysis to generate a regional subset of data which I compare with datasets describing co-distributed taxa, SDC species and sympatric sister species. Results A weak negative relationship between FST and dispersal potential is implied by aggregate global analysis (0.1 ≤ R2 ≤ 0.29). In contrast, regional datasets of co-distributed species show strong correlation between FST and dispersal potential (0.78 ≤ R2 ≤ 0.85). Comparisons between SDC and sympatric sister species consistently evince higher gene flow in species with higher dispersal potential. Main conclusions Ad hoc aggregate analyses can be compromised by multiple sources of error. Comparisons of SDC taxa, including sympatric sister species, adapt the experimental scientific method to natural situations, providing robust and repeatable tests of phylogeographic hypotheses and accurate estimates of effect sizes. To make strong inferences about phylogeography we should seek out the hidden wealth of natural experiments that provide particularly clear opportunities to study the factors that influence patterns of biodiversity. The SDC framework enables independent tests of existing hypotheses, integration of independent studies in meta-analyses across taxa and regions, and identification of general trends versus location- or taxon-specific phenomena. Coupled with advances in statistical phylogeography, species distribution modelling and palaeoecology, the SDC framework bridges the long-standing gap between observational and experimental sciences. [ABSTRACT FROM AUTHOR]

Aim We still have limited understanding of the contingent and deterministic factors that have fostered the evolutionary success of some species lineages over others. We investigated how the interplay of intercontinental migration and key innovations promoted diversification of the genus Androsace. Location Mountain ranges and cold steppes of the Northern Hemisphere. Methods We reconstructed ancestral biogeographical ranges at regional and continental scales by means of a dispersal-extinction-cladogenesis analysis using dated Bayesian phylogenies and contrasting two migration scenarios. Based on diversification analyses under two frameworks, we tested the influence of life form on speciation rates and whether diversification has been diversity-dependent. Results We found that three radiations occurred in this genus, at different periods and on different continents, and that life form played a critical role in the history of Androsace. Short-lived ancestors first facilitated the expansion of the genus' range from Asia to Europe, while cushions, which appeared independently in Asia and Europe, enhanced species diversification in alpine regions. One long-distance dispersal event from Europe to North America led to the diversification of the nested genus Douglasia. We found support for a model in which speciation of the North American- European clade is diversity-dependent and close to its carrying capacity, and that the diversification dynamics of the North American subclade are uncoupled from this and follow a pure birth process. Main conclusions The contingency of past biogeographical connections combined with the evolutionary determinism of convergent key innovations may have led to replicated radiations of Androsace in three mountain regions of the world. The repeated emergence of the cushion life form was a convergent key innovation that fostered radiation into alpine habitats. Given the large ecological similarity of Androsace species, allopatry may have been the main mode of speciation. [ABSTRACT FROM AUTHOR]

Aim Macroecological patterns have mainly been depicted as atemporal, with existing research covering only short time periods. One fundamental pattern in macroecology is the interspecific relationship between local abundance and regional range size, which is generally considered to be positively linear. Here, we examine structural details of the relationship between abundance and range size in cyclic populations of small mammals and its long-term temporal variation. Location Finland. Methods We analysed 39 years of trapping data of Rodentia and Soricomorpha collected in field and forest habitats across Finland. Abundance was measured as the mean population density of individuals, and range size as the number of grid cells occupied and sample occupancy. Data were analysed using linear mixed models. Results The relationship between mean local density and sample occupancy was generally linear, whereas the relationship between density and the number of grid cells occupied was generally curvilinear (U-shaped) for both habitats, being negative for species with small range sizes and positive for species with larger ranges. The curvilinearity was temporally consistent in both habitats. Main conclusions The interspecific relationship between abundance and range size varies depending on how range size is measured. Interestingly, the curvilinear relationship between density and range size was found in an assemblage in which many of the species have fluctuating (cyclic) populations. Future research on abundance-range size relationships should focus on long-term temporal variation to better understand the underlying mechanisms and to develop macroecological theory. [ABSTRACT FROM AUTHOR]

Understanding the processes responsible for macro-scale spatial and temporal phenological patterns is a critical step in developing predictive phenological models. While phenological responses may involve the integration of multiple environmental cues, the spring phenology of many plant and animal species appears to be especially sensitive to temperature., As a result of the success of citizen science schemes in mobilizing amateur naturalists, for some parts of the world, there now exist extensive data sets of phenological timings, spanning many species, locations and years. In macroecology, two types of models - time windows and growing degree-days - are widely used to predict phenology on the basis of temperature., Here, we compare the performance of the two methods in predicting spatiotemporal variation in the timing of Quercus robur first leafing. The methods agree on the time at which leafing becomes sensitive to temperature and provide weak support for a delay in initiation of thermal sensitivity with increasing latitude due to a day-length requirement. Both methods explain c. 50% of the variation in first dates and identify plasticity, rather than local adaptation, as the major cause of spatial covariation between temperature and phenology. For a 1°C rise in spring temperatures we predict that a plastic response of first leafing will give rise to an advance of about seven days., Synthesis: Time-window and growing degree-day methods provide remarkably congruent insights into the processes underpinning geographic variation in Quercus robur first leafing dates. We find that a spatially invariant plastic response to temperature dominates spatiotemporal phenological variation, which means that it may be reasonable to substitute space for time to project how this species will respond to climate change. This study demonstrates the contribution that top-down macroecological approaches can make to our understanding of the processes that give rise to intraspecific phenological variation. [ABSTRACT FROM AUTHOR]

Aim Using sequences of complete mitochondrial genomes, our aims were: (1) to investigate the matrilineal phylogeographical structure, migration patterns and lineage coalescence times in a large, continuous population of brown bears ( Ursus arctos); and (2) to develop a novel spatial genetic method to identify migration corridors and barriers. Location North-western Eurasia: from eastern European Russia to the Baltic Sea. Methods We sequenced the complete mitochondrial genomes of 95 brown bears. The phylogeographical resolution of complete genomes was compared to that derived from subsets of the genome, including the most commonly used shorter sequences. We conducted network and Bayesian phylogeographical analyses and developed a novel, spatially explicit, individual-based approach (called DResD) for identifying migration corridors and barriers. Results Analysis of mitogenome sequences revealed five haplogroups, specific to particular geographical areas, exhibiting far greater resolving power than shorter sequences. Estimated coalescence times for the haplogroups ranged from 7.7 to 15.2 ka, suggesting that their divergence took place after the last glaciation. We found several migration trends, including a large westward migration from eastern European Russia towards Finland. We also found evidence of a potential barrier and a migration corridor in the south-west of the study area. Main conclusions The use of complete mitochondrial genomes from a brown bear population in north-western Eurasia allowed us to identify phylogeographical structure, signatures of demographic history and spatial processes that had not previously been detected using shorter sequences. These findings have implications for studies on other species and populations, especially those exhibiting low mt DNA diversity. The relatively recent divergence estimates for haplogroups highlight the significance not only of the last glaciation but also of climatic fluctuations during the post-glacial period for the divergence of mammal populations in Europe. Our spatial genetic method represents a new tool for the analysis of genetic data in a geographical context and is applicable to any data that yield genetic distance matrices, including microsatellites, amplified fragment length polymorphisms ( AFLPs) and single-nucleotide polymorphisms ( SNPs). [ABSTRACT FROM AUTHOR]

A major aim of island biogeography has been to describe general patterns of species richness across islands and to identify the processes responsible. Data are often collected across many islands; with larger datasets providing increased statistical power and more accurate parameter estimates. However, there is often structure in observational data, violating an assumption of linear models that each datum is independent. In island biogeography this structure may take the form of an island, archipelago or taxon being represented by multiple data points. We survey recent papers in this field and find that these forms of non-independence are a common feature. Most authors addressed this problem by conducting separate analyses for each archipelago, taxon or combination of the two, but a better tool for dealing with non-independence and structure in data, the mixed model, already exists. We demonstrate the advantages of a mixed model approach by applying it to a well-known dataset that spans 134 observations of single island endemic (SIE) richness across 39 islands, four archipelagos and four taxa. Taking island area and age into account, SIE richness varies substantially more among archipelagos than it does among islands or taxa. We find that SIE richness rises with island age on the Azores and Galapagos, while on the Canaries and Hawaii SIE richness initially rises with age but later declines on older islands. Our analyses demonstrate three advantages to island biogeography of applying a mixed modelling approach: 1) structure in the data is controlled for; 2) the variance among islands, archipelagos and taxa is estimated; 3) all the data can be included in a single model, making it possible to test whether trends are general across all archipelagos and taxa or are idiosyncratic. [ABSTRACT FROM AUTHOR]

Island taxa often attain forms outside the range achieved by mainland relatives. Body size evolution of vertebrates on islands has therefore received much attention, with two seemingly conflicting patterns thought to prevail: (1) islands harbour animals of extreme size, and (2) islands promote evolution towards medium body size ('the island rule'). We test both hypotheses using body size distributions of mammal, lizard and bird species. World-wide. We assembled body size and insularity datasets for the world's lizards, birds and mammals. We compared the frequencies with which the largest or smallest member of a group is insular with the frequencies expected if insularity is randomly assigned within groups. We tested whether size extremes on islands considered across mammalian phylogeny depart from a null expectation under a Brownian motion model. We tested the island rule by comparing insular and mainland members of (1) a taxonomic level and (2) mammalian sister species, to determine if large insular animals tend to evolve smaller body sizes while small ones evolve larger sizes. The smallest species in a taxon (order, family or genus) are insular no more often than would be expected by chance in all groups. The largest species within lizard families and bird genera (but no other taxonomic levels) are insular more often than expected. The incidence of extreme sizes in insular mammals never departs from the null, except among extant genera, where gigantism is marginally less common than expected under a Brownian motion null. Mammals follow the island rule at the genus level and when comparing sister species and clades. This appears to be driven mainly by insular dwarfing in large-bodied lineages. A similar pattern in birds is apparent for species within orders. However, lizards follow the converse pattern. The popular misconception that islands have more than their fair share of size extremes may stem from a greater tendency to notice gigantism and dwarfism when they occur on islands. There is compelling evidence for insular dwarfing in large mammals, but not in other taxa, and little evidence for the second component of the island rule - gigantism in small-bodied taxa. [ABSTRACT FROM AUTHOR]

Aim (1) To describe the species–area relationships among communities of Plasmodium and Haemoproteus parasites in different island populations of the same host genus (Aves: Zosterops). (2) To compare distance–decay relationships (turnover) between parasite communities and those with potential avian and dipteran hosts, which differ with respect to their movement and potential to disperse parasite species over large distances. Location Two archipelagos in the south-west Pacific, Vanuatu and New Caledonia ( c. 250 km west of Vanuatu) and its Loyalty Islands, with samples collected from a total of 16 islands of varying sizes (328–16,648 km2). Methods We characterized parasite diversity and distribution via polymerase chain reaction (PCR) from avian ( Zosterops) blood samples. Bayesian methods were used to reconstruct the parasite phylogeny. In accordance with recent molecular evidence, we treat distinct mitochondrial DNA lineages as equivalent to species in this study. Path analysis and parasite lineage accumulation curves were used to assess the confounding effect of inadequate sampling on the estimation of parasite richness. Species–area and species–distance relationships were assessed using linear regression: distance–decay relationships were assessed using Mantel tests. Results Birds and mosquito species and Plasmodium lineages exhibited significant species–area relationships. However, Plasmodium lineages showed the weakest ‘species–area’ relationship; no relationship was found for Haemoproteus lineages. Avian species richness influenced parasite lineage richness more than mosquito species richness did. Within individual avian host species, the species–area relationship of parasites showed differing patterns. Path analysis indicated that sampling effort was unlikely to have a confounding effect on parasite richness. Distance from mainland (isolation effect) showed no effect on parasite richness. Community similarity decayed significantly with distance for avifauna, mosquito fauna and Plasmodium lineages but not for Haemoproteus lineages. Main conclusions Plasmodium lineages and mosquito species fit the power-law model with steeper slopes than found for the avian hosts. The lack of species–distance relationship in parasites suggests that other factors, such as the competence of specific vectors and habitat features, may be more important than distance. The decay in similarity with distance suggests that the sampled Plasmodium lineages and their potential hosts were not randomly distributed, but rather exhibited spatially predictable patterns. We discuss these results in the context of the effects that parasite generality may have on distribution patterns. [ABSTRACT FROM AUTHOR]

1. Dispersal is one of the key ecological parameters but it is very difficult to quantify directly. As a consequence, empirical studies often ignore dispersal or use indirect measures. 2. Ringing data have previously been used to estimate the natal dispersal distances of 47 British passerine bird species. This provides an excellent opportunity to examine the potential of various indirect measures to predict natal dispersal distances in British birds. 3. We use a phylogenetic comparative framework and single- and multipredictor models including ecomorphological, behavioural or ecological traits to predict natal dispersal distance. 4. A multipredictor model that includes Kipp's distance (a measure of wing tip length), bill depth and tail graduation explains 45% of the interspecific variation in natal dispersal distance. These morphological characters all relate to aerodynamics with stronger flyers dispersing further. 5. However, an index of migration is a strong (but less informative) correlate of dispersal distance and Kipp's distance and bill depth are strong correlates of migration. Thus, we cannot disentangle whether these ecomorphological traits influence dispersal distance directly or whether the relationship between ecomorphology and dispersal is mediated through migratory behaviour. 6. Notwithstanding uncertainties regarding the causal links between dispersal distance and wing morphology, we suggest that two ecomorphological traits, Kipp's distance and bill depth, may provide a useful surrogate. [ABSTRACT FROM AUTHOR]

The article focuses on the study about the island rule that describes a tendency for larger animals to get smaller on islands, and for smaller ones to get larger, as well as the reduced major axis regrassion. Intricately, the rule has been assessed by comparing matched pairs of species or populations on islands with their presumed continental source. Results show that the variance in body size among island species is roughly the same as the variance in body size among continental species. Analysis of the study also implies that there is a general tendency for body sizes to converge on some intermediate value on islands.