Evolution of wing shape in geometrid moths: phylogenetic effects dominate over ecology.

Locomotory performance is an important determinant of fitness in most animals, including flying insects. Strong selective pressures on wing morphology are therefore expected. Previous studies on wing shape in Lepidoptera have found some support for hypotheses relating wing shape to environment-specific selective pressures on aerodynamic performance. Here, we present a phylogenetic comparative study on wing shape in the lepidopteran family Geometridae, covering 374 species of the northern European fauna. We focused on 11 wing traits including aspect ratio, wing roundness, and the pointedness of the apex, as well as the ratio of forewing and hindwing areas. All measures were taken from images available on the internet, using a combination of tools available in Fiji software and R. We found that wing shape demonstrates a phylogenetically conservative pattern of evolution in Geometridae, showing similar or stronger phylogenetic signal than many of its potential predictors. Several wing traits showed statistically significant associations with predictors such as body size, phenology, and preference for forest habitats. Overall, however, all of these associations remained notably weak, with no wing shape being excluded for any value of the predictors, including body size. We conclude that, in geometrids, wing traits do not readily respond to selective pressures optimizing aerodynamic performance of the moths in different environments. Selection on wing shape may nevertheless operate through other functions of the wings, with the effectiveness of crypsis at rest being a promising candidate for further studies.

Modeling forest landscape futures: Full scale simulation of realistic socioeconomic scenarios in Estonia.

For political and administrative governance of land-use decisions, high-resolution and reliable spatial models are required over large areas and for various time horizons. We present a process-centered simulation model 'NextStand' (a forest landscape model, FLM) and its R-script, which predicts regional forest characteristics at a forest stand resolution. The model uses whole area stand data and is optimized for realistic iterative timber harvesting decisions, based on stand compositions (developing over time) and locations. We used the model for simulating spatial predictions of the Estonian forests in North Europe (2.3 Mha, about 2 M stands); the decisions were parameterized by land ownership, protection regimes, and rules of clear-cut harvesting. We illustrate the model application as a potential broad-scale Decision Support Tool by predicting how the forest age composition, placement of clear-cut areas, and connectivity of old stands will develop until the year 2050 under future scenarios. The country-scale outputs had a generally low within-scenario variance, which enabled to estimate some main land-use effects and uncertainties at small computing efforts. In forestry terms, we show that a continuation of recent intensive forest management trends will produce a decline of the national timber supplies in Estonia, which greatly varies among ownership types. In a conservation perspective, the current level of 13% forest area strictly protected can maintain an overall area of old forests by 2050, but their isolation is a problem for biodiversity conservation. The behavior of low-intensity forest management units (owners) and strict governance of clear-cut harvesting rules emerged as key questions for regional forest sustainability. Our study confirms that high-resolution modeling of future spatial composition of forest land is feasible when one can (i) delineate predictable spatial units of transformation (including management) and (ii) capture their variability of temporal change with simple ecological and socioeconomic (including human decision-making) variables.

Early-life food stress hits females harder than males in insects: A meta-analysis of sex differences in environmental sensitivity.

Fitness consequences of early-life environmental conditions are often sex-specific, but corresponding evidence for invertebrates remains inconclusive. Here, we use meta-analysis to evaluate sex-specific sensitivity to larval nutritional conditions in insects. Using literature-derived data for 85 species with broad phylogenetic and ecological coverage, we show that females are generally more sensitive to food stress than males. Stressful nutritional conditions during larval development typically lead to female-biased mortality and thus increasingly male-biased sex ratios of emerging adults. We further demonstrate that the general trend of higher sensitivity to food stress in females can primarily be attributed to their typically larger body size in insects and hence higher energy needs during development. By contrast, there is no consistent evidence of sex-biased sensitivity in sexually size-monomorphic species. Drawing conclusions regarding sex-biased sensitivity in species with male-biased size dimorphism remains to wait for the accumulation of relevant data. Our results suggest that environmental conditions leading to elevated juvenile mortality may potentially affect the performance of insect populations further by reducing the proportion of females among individuals reaching reproductive age. Accounting for sex-biased mortality is therefore essential to understanding the dynamics and demography of insect populations, not least importantly in the context of ongoing insect declines.

Effects of air humidity and soil moisture on secondary metabolites in the leaves and roots of Betula pendula of different competitive status.

Plant secondary metabolites (PSMs) defend plants against abiotic stresses, including those caused by climate change and against biotic stresses, such as herbivory and competition. There is a trade-off between allocating available carbon to growth and defence in stressful environments. However, our knowledge about trade-off is limited, especially when abiotic and biotic stresses co-occur. We aimed to understand the combined effect of increasing precipitation and humidity, the tree's competitive status, and canopy position on leaf secondary metabolites (LSMs) and fine root secondary metabolites (RSMs) in Betula pendula. We sampled 8-year-old B. pendula trees growing in the free air humidity manipulation (FAHM) experimental site, where treatments included elevated relative air humidity and elevated soil moisture. A high-performance liquid chromatography-quadrupole-time of flight mass spectrometer (HPLC-qTOF-MS) was used to analyse secondary metabolites. Our results showed accumulation of LSM depends on the canopy position and competitive status. Flavonoids (FLA), dihydroxybenzoic acids (HBA), jasmonates (JA) and terpene glucosides (TG) were higher in the upper canopy, and FLA, monoaryl compounds (MAR) and sesquiterpenoids (ST) were higher in dominant trees. The FAHM treatments had a more distinct effect on RSM than on LSM. The RSMs were lower in elevated air humidity and soil moisture conditions than in control conditions. The RSM content depended on the competitive status and was higher in suppressed trees. Our study suggests that young B. pendula will allocate similar amounts of carbon to constitutive chemical leaf defence, but a lower amount to root defence (per fine root biomass) under higher humidity.

Limited sex differences in plastic responses suggest evolutionary conservatism of thermal reaction norms: A meta-analysis in insects.

Temperature has a profound effect on the growth and development of ectothermic animals. However, the extent to which ecologically driven selection pressures can adjust thermal plastic responses in growth schedules is not well understood. Comparing temperature-induced plastic responses between sexes provides a promising but underexploited approach to evaluating the evolvability of thermal reaction norms: males and females share largely the same genes and immature environments but typically experience different ecological selection pressures. We proceed from the idea that substantial sex differences in plastic responses could be interpreted as resulting from sex-specific life-history optimization, whereas similarity among the sexes should rather be seen as evidence of an essential role of physiological constraints. In this study, we performed a meta-analysis of sex-specific thermal responses in insect development times, using data on 161 species with comprehensive phylogenetic and ecological coverage. As a reference for judging the magnitude of sex specificity in thermal plasticity, we compared the magnitude of sex differences in plastic responses to temperature with those in response to diet. We show that sex-specific responses of development times to temperature variation are broadly similar. We also found no strong evidence for sex specificity in thermal responses to depend on the magnitude or direction of sex differences in development time. Sex differences in temperature-induced plastic responses were systematically less pronounced than sex differences in responses induced by variations in larval diet. Our results point to the existence of substantial constraints on the evolvability of thermal reaction norms in insects as the most likely explanation. If confirmed, the low evolvability of thermal response is an essential aspect to consider in predicting evolutionary responses to climate warming.

Assessing cumulative impacts of human-induced pressures on reef and sandbank habitats and associated biotopes in the northeastern Baltic Sea.

Marine ecosystems are impacted by multiple individual and combined anthropogenic pressures. We used meta-analysis and data-driven PlanWise4Blue decision support tool to predict individual and combined impacts of wind park development, nutrient loading, and invasive species on vulnerable reef and sandbank habitats and associated species-specific biotopes in the northeastern Baltic Sea. Many impacts were not statistically significant due to large between-study variance in effect sizes. Wind park development is predicted to have less impact than nutrient loading and invasive species. Predicted impacts varied greatly among larger-scale habitats versus smaller-scale biotopes with impacts being generally stronger at small scale. Excessive nutrient loading damages algae-based biotopes, the presence of nonnative species has substantial negative impacts on larger-scale reef and sandbank habitats. The results showed that a 25 % reduction of nutrient loading improves all examined benthic habitats, whereas nonnative species, which cannot be removed from ecosystems, pose a significant threat to these habitats.

Assessing the potential for sea-based macroalgae cultivation and its application for nutrient removal in the Baltic Sea.

Marine eutrophication is a pervasive and growing threat to global sustainability. Macroalgal cultivation is a promising circular economy solution to achieve nutrient reduction and food security. However, the location of production hotspots is not well known. In this paper the production potential of macroalgae of high commercial value was predicted across the Baltic Sea region. In addition, the nutrient limitation within and adjacent to macroalgal farms was investigated to suggest optimal site-specific configuration of farms. The production potential of Saccharina latissima was largely driven by salinity and the highest production yields are expected in the westernmost Baltic Sea areas where salinity is >23. The direct and interactive effects of light availability, temperature, salinity and nutrient concentrations regulated the predicted changes in the production of Ulva intestinalis and Fucus vesiculosus. The western and southern Baltic Sea exhibited the highest farming potential for these species, with promising areas also in the eastern Baltic Sea. Macroalgal farming did not induce significant nutrient limitation. The expected spatial propagation of nutrient limitation caused by macroalgal farming was less than 100-250 m. Higher propagation distances were found in areas of low nutrient and low water exchange (e.g. offshore areas in the Baltic Proper) and smaller distances in areas of high nutrient and high water exchange (e.g. western Baltic Sea and Gulf of Riga). The generated maps provide the most sought-after input to support blue growth initiatives that foster the sustainable development of macroalgal cultivation and reduction of in situ nutrient loads in the Baltic Sea.

Blueprint for the ideal microplastic effect study: Critical issues of current experimental approaches and envisioning a path forward.

This article presents a novel conceptual blueprint for an 'ideal', i.e., ecologically relevant, microplastic effect study. The blueprint considers how microplastics should be characterized and applied in laboratory experiments, and how biological responses should be measured to assure unbiased data that reliably reflect the effects of microplastics on aquatic biota. This 'ideal' experiment, although practically unachievable, serves as a backdrop to improve specific aspects of experimental research on microplastic effects. In addition, a systematic and quantitative literature review identified and quantified departures of published experiments from the proposed 'ideal' design. These departures are related mainly to the experimental design of microplastic effect studies failing to mimic natural environments, and experiments with limited potential to be scaled-up to ecosystem level. To produce a valid and generalizable assessment of the effect of microplastics on biota, a quantitative meta-analysis was performed that incorporated the departure of studies from the 'ideal' experiment (a measure of experimental quality) and inverse variance (a measure of the study precision) as weighting coefficients. Greater weights were assigned to experiments with higher quality and/or with lower variance in the response variables. This double-weighting captures jointly the technical quality, ecological relevance and precision of estimates provided in each study. The blueprint and associated meta-analysis provide an improved baseline for the design of ecologically relevant and technically sound experiments to understand the effects of microplastics on single species, populations and, ultimately, entire ecosystems.

Climate and Competitive Status Modulate the Variation in Secondary Metabolites More in Leaves Than in Fine Roots of Betula pendula.

Plant secondary metabolites have many important functions; they also determine the productivity and resilience of trees under climate change. The effects of environmental factors on secondary metabolites are much better understood in above-ground than in below-ground part of the tree. Competition is a crucial biotic stress factor, but little is known about the interaction effect of climate and competition on the secondary chemistry of trees. Moreover, competition effect is usually overlooked when analyzing the sources of variation in the secondary chemistry. Our aim was to clarify the effects of competitive status, within-crown light environment, and climate on the secondary chemistry of silver birch (Betula pendula Roth). We sampled leaves (from upper and lower crown) and fine roots from competitively dominant and suppressed B. pendula trees in plantations along a latitudinal gradient (56-67° N) in Fennoscandia, with mean annual temperature (MAT) range: -1 to 8°C. Secondary metabolites in leaves (SML) and fine roots (SMFR) were determined with an HPLC-qTOF mass spectrometer. We found that SML content increased significantly with MAT. The effect of competitive stress on SML strengthened in colder climates (MAT<4°C). Competition and shade initiated a few similar responses in SML. SMFR varied less with MAT. Suppressed trees allocated relatively more resources to SML in warmer climates and to SMFR in colder ones. Our study revealed that the content and profile of secondary metabolites (mostly phenolic defense compounds and growth regulators) in leaves of B. pendula varied with climate and reflected the trees' defense requirements against herbivory, exposure to irradiance, and competitive status (resource supply). The metabolic profile of fine roots reflected, besides defense requirements, also different below-ground competition strategies in warmer and colder climates. An increase in carbon assimilation to secondary compounds can be expected at northern latitudes due to climate change.

The initial overreaction of carbon cycle to elevated atmospheric humidity levels off over time - a FAHM study in a young birch forest.

Ecosystem responses to climate change are mainly predicted based on short-term studies. However, the first response can be a temporary overreaction, different from the later response of the more acclimated ecosystem. The current paper is a follow-up study of our previous article, where the effect of elevated atmospheric humidity on forest ecosystem carbon (C) balance was studied in a young silver birch (Betula pendula Roth) forest after two years of humidification. Here, we present the C balance of the same forest measured two years later when humidification treatment had been performed for four years. We revealed that the higher C sequestration capacity of the humidified birch forest ecosystem was an initial overreaction, which levelled off after four years of humidification, when the ecosystem became more acclimated to wetter conditions. Understorey production reacted rapidly and strongly by increasing belowground production more than twofold, but this reaction ceased after four years of humidification treatment. Trees responded to a lesser extent, and the initially decreased aboveground growth was recovered after four years of humidification, when the biomass allocation to tree fine-roots was increased. Our results showed that at early forest age, understorey plant production dominated in the whole ecosystem C sequestration capacity. But in the later stage, the most important C sink was biomass production of birches, and since the tree biomass production no longer differed between the treatments, C sequestration of the whole ecosystem did not differ either. The findings confirm that a preliminary reaction of an ecosystem can be different from the later response, which needs to be taken into account when prognosing the climate change consequences for carbon sequestration.

Why do males emerge before females? Sexual size dimorphism drives sexual bimaturism in insects.

Conspecific females and males often follow different development trajectories which leads to sex differences in age at maturity (sexual bimaturism, SBM). Whether SBM is typically selected for per se (direct selection hypothesis) or merely represents a side-effect of other sex-related adaptations (indirect selection hypothesis) is, however, still an open question. Substantial interspecific variation in the direction and degree of SBM, both in invertebrates and vertebrates, calls for multi-species studies to understand the relative importance of its evolutionary drivers. Here we use two complementary approaches to evaluate the evolutionary basis of SBM in insects. For this purpose, we assembled an extensive literature-derived data set of sex-specific development times and body sizes for a taxonomically and ecologically wide range of species. We use these data in a meta-analytic framework to evaluate support for the direct and indirect selection hypotheses. Our results confirm that protandry - males emerging as adults before females - is the prevailing form of SBM in insects. Nevertheless, protandry is not as ubiquitous as often presumed: females emerged before males (= protogyny) in about 36% of the 192 species for which we had data. Moreover, in a considerable proportion of species, the sex difference in the timing of adult emergence was negligible. In search for the evolutionary basis of SBM, we found stronger support for the hypothesis that explains SBM by indirect selection. First, across species, the direction and degree of SBM appeared to be positively associated with the direction and degree of sexual size dimorphism (SSD). This is consistent with the view that SBM is a correlative by-product of evolution towards sexually dimorphic body sizes. Second, within protandrous species, the degree of protandry typically increased with plastic increase in development time, with females prolonging their development more than males in unfavourable conditions. This pattern is in conflict with the direct selection hypothesis, which predicts the degree of protandry to be insensitive to the quality of the juvenile environment. These converging lines of evidence support the idea that, in insects, SBM is generally a by-product of SSD rather than a result of selection on the two sexes to mature at different times. It appears plausible that selective pressures on maturation time per se generally cannot compete with viability- and fecundity-mediated selection on insect body sizes. Nevertheless, exceptions certainly exist: there are undeniable cases of SBM where this trait has evolved in response to direct selection. In such cases, either the advantage of sex difference in maturation time must have been particularly large, or fitness effects of body size have been unusually weak.

Predawn leaf conductance depends on previous day irradiance but is not related to growth in aspen saplings grown under artificially manipulated air humidity.

Recent studies have suggested that predawn stomatal opening may enhance early-morning photosynthesis (A) and improve the relative growth rate of trees. However, the causality between night-time stomatal conductance, A, and tree growth is disputable because stomatal opening in darkness can be mediated by previous day photosynthate loads and might be a consequence of growth-related processes like dark respiration (R). To identify linkages between night-time leaf conductance (gl_night), A, R, and tree growth, we conducted an experiment in hybrid aspen saplings grown under different air relative humidity (RH) conditions and previous day irradiance level (IR_pday). Predawn leaf conductance (gl_predawn) depended on RH, IR_pday and R (P < 0.05), whereas early-morning gross A (Agross_PAR500) depended on IR_pday and gl_predawn (P < 0.001). Daytime net A was positively related to Agross_PAR500 and leaf [N] (P < 0.05). Tree diameter and height increment correlated positively with gl at the beginning and middle of the night (P < 0.05) but not before dawn. Although our results demonstrate that gl_night was related to tree growth, the relationship was not determined by R. The linkage between gl_predawn and Agross_PAR500 was modified by IR_pday, indicating that daily CO2 assimilation probably provides feedback for stomatal opening before dawn.

Trans-Atlantic Distribution and Introgression as Inferred from Single Nucleotide Polymorphism: Mussels Mytilus and Environmental Factors.

Large-scale climate changes influence the geographic distribution of biodiversity. Many taxa have been reported to extend or reduce their geographic range, move poleward or displace other species. However, for closely related species that can hybridize in the natural environment, displacement is not the only effect of changes of environmental variables. Another option is subtler, hidden expansion, which can be found using genetic methods only. The marine blue mussels Mytilus are known to change their geographic distribution despite being sessile animals. In addition to natural dissemination at larval phase-enhanced by intentional or accidental introductions and rafting-they can spread through hybridization and introgression with local congeners, which can create mixed populations sustaining in environmental conditions that are marginal for pure taxa. The Mytilus species have a wide distribution in coastal regions of the Northern and Southern Hemisphere. In this study, we investigated the inter-regional genetic differentiation of the Mytilus species complex at 53 locations in the North Atlantic and adjacent Arctic waters and linked this genetic variability to key local environmental drivers. Of seventy-nine candidate single nucleotide polymorphisms (SNPs), all samples were successfully genotyped with a subset of 54 SNPs. There was a clear interregional separation of Mytilus species. However, all three Mytilus species hybridized in the contact area and created hybrid zones with mixed populations. Boosted regression trees (BRT) models showed that inter-regional variability was important in many allele models but did not prevail over variability in local environmental factors. Local environmental variables described over 40% of variability in about 30% of the allele frequencies of Mytilus spp. For the 30% of alleles, variability in their frequencies was only weakly coupled with local environmental conditions. For most studied alleles the linkages between environmental drivers and the genetic variability of Mytilus spp. were random in respect to "coding" and "non-coding" regions. An analysis of the subset of data involving functional genes only showed that two SNPs at Hsp70 and ATPase genes correlated with environmental variables. Total predictive ability of the highest performing models (r2 between 0.550 and 0.801) were for alleles that discriminated most effectively M.trossulus from M.edulis and M.galloprovincialis, whereas the best performing allele model (BM101A) did the best at discriminating M.galloprovincialis from M. edulis and M.trossulus. Among the local environmental variables, salinity, water temperature, ice cover and chlorophyll a concentration were by far the greatest predictors, but their predictive performance varied among different allele models. In most cases changes in the allele frequencies along these environmental gradients were abrupt and occurred at a very narrow range of environmental variables. In general, regions of change in allele frequencies for M.trossulus occurred at 8-11 psu, 0-10 C, 60%-70% of ice cover and 0-2 mg m-3 of chlorophyll a, M. edulis at 8-11 and 30-35 psu, 10-14 C and 60%-70% of ice cover and for M.galloprovincialis at 30-35 psu, 14-20 C.

Cleaning up seas using blue growth initiatives: Mussel farming for eutrophication control in the Baltic Sea.

Eutrophication is a serious threat to aquatic ecosystems globally with pronounced negative effects in the Baltic and other semi-enclosed estuaries and regional seas, where algal growth associated with excess nutrients causes widespread oxygen free "dead zones" and other threats to sustainability. Decades of policy initiatives to reduce external (land-based and atmospheric) nutrient loads have so far failed to control Baltic Sea eutrophication, which is compounded by significant internal release of legacy phosphorus (P) and biological nitrogen (N) fixation. Farming and harvesting of the native mussel species (Mytilus edulis/trossulus) is a promising internal measure for eutrophication control in the brackish Baltic Sea. Mussels from the more saline outer Baltic had higher N and P content than those from either the inner or central Baltic. Despite their relatively low nutrient content, harvesting farmed mussels from the central Baltic can be a cost-effective complement to land-based measures needed to reach eutrophication status targets and is an important contributor to circularity. Cost effectiveness of nutrient removal is more dependent on farm type than mussel nutrient content, suggesting the need for additional development of farm technology. Furthermore, current regulations are not sufficiently conducive to implementation of internal measures, and may constitute a bottleneck for reaching eutrophication status targets in the Baltic Sea and elsewhere.

Quantifying the effects of species traits on predation risk in nature: A comparative study of butterfly wing damage.

Evading predators is a fundamental aspect of the ecology and evolution of all prey animals. In studying the influence of prey traits on predation risk, previous researchers have shown that crypsis reduces attack rates on resting prey, predation risk increases with increased prey activity, and rapid locomotion reduces attack rates and increases chances of surviving predator attacks. However, evidence for these conclusions is nearly always based on observations of selected species under artificial conditions. In nature, it remains unclear how defensive traits such as crypsis, activity levels and speed influence realized predation risk across species in a community. Whereas direct observations of predator-prey interactions in nature are rare, insight can be gained by quantifying bodily damage caused by failed predator attacks. We quantified how butterfly species traits affect predation risk in nature by determining how defensive traits correlate with wing damage caused by failed predation attempts, thereby providing the first robust multi-species comparative analysis of predator-induced bodily damage in wild animals. For 34 species of fruit-feeding butterflies in an African forest, we recorded wing damage and quantified crypsis, activity levels and flight speed. We then tested for correlations between damage parameters and species traits using comparative methods that account for measurement error. We detected considerable differences in the extent, location and symmetry of wing surface loss among species, with smaller differences between sexes. We found that males (but not females) of species that flew faster had substantially less wing surface loss. However, we found no correlation between cryptic coloration and symmetrical wing surface loss across species. In species in which males appeared to be more active than females, males had a lower proportion of symmetrical wing surface loss than females. Our results provide evidence that activity greatly influences the probability of attacks and that flying rapidly is effective for escaping pursuing predators in the wild, but we did not find evidence that cryptic species are less likely to be attacked while at rest.

Publisher Correction: Contrasting latitudinal patterns in phylogenetic diversity between woody and herbaceous communities.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

Contrasting latitudinal patterns in phylogenetic diversity between woody and herbaceous communities.

Although many studies have shown that species richness decreases from low to high latitudes (the Latitudinal Diversity Gradient), little is known about the relationship between latitude and phylogenetic diversity. Here we examine global latitudinal patterns of phylogenetic diversity using a dataset of 459 woody and 589 herbaceous plant communities. We analysed the relationships between community phylogenetic diversity, latitude, biogeographic realm and vegetation type. Using the most recent global megaphylogeny for seed plants and the standardised effect sizes of the phylogenetic diversity metrics 'mean pairwise distance' (SESmpd) and 'mean nearest taxon distance' (SESmntd), we found that species were more closely-related at low latitudes in woody communities. In herbaceous communities, species were more closely-related at high latitudes than at intermediate latitudes, and the strength of this effect depended on biogeographic realm and vegetation type. Possible causes of this difference are contrasting patterns of speciation and dispersal. Most woody lineages evolved in the tropics, with many gymnosperms but few angiosperms adapting to high latitudes. In contrast, the recent evolution of herbaceous lineages such as grasses in young habitat types may drive coexistence of closely-related species at high latitudes. Our results show that high species richness commonly observed at low latitudes is not associated with high phylogenetic diversity.

Linking atmospheric, terrestrial and aquatic environments: Regime shifts in the Estonian climate over the past 50 years.

Climate change in recent decades has been identified as a significant threat to natural environments and human wellbeing. This is because some of the contemporary changes to climate are abrupt and result in persistent changes in the state of natural systems; so called regime shifts (RS). This study aimed to detect and analyse the timing and strength of RS in Estonian climate at the half-century scale (1966-2013). We demonstrate that the extensive winter warming of the Northern Hemisphere in the late 1980s was represented in atmospheric, terrestrial, freshwater and marine systems to an extent not observed before or after the event within the studied time series. In 1989, abiotic variables displayed statistically significant regime shifts in atmospheric, river and marine systems, but not in lake and bog systems. This was followed by regime shifts in the biotic time series of bogs and marine ecosystems in 1990. However, many biotic time series lacked regime shifts, or the shifts were uncoupled from large-scale atmospheric circulation. We suggest that the latter is possibly due to complex and temporally variable interactions between abiotic and biotic elements with ecosystem properties buffering biotic responses to climate change signals, as well as being affected by concurrent anthropogenic impacts on natural environments.

A review of urban impacts on avian life-history evolution: Does city living lead to slower pace of life?

The concept of a pace-of-life syndrome describes inter- and intraspecific variation in several life-history traits along a slow-to-fast pace-of-life continuum, with long lifespans, low reproductive and metabolic rates, and elevated somatic defences at the slow end of the continuum and the opposite traits at the fast end. Pace-of-life can vary in relation to local environmental conditions (e.g. latitude, altitude), and here we propose that this variation may also occur along an anthropogenically modified environmental gradient. Based on a body of literature supporting the idea that city birds have longer lifespans, we predict that urban birds have a slower pace-of-life compared to rural birds and thus invest more in self maintenance and less in annual reproduction. Our statistical meta-analysis of two key traits related to pace-of-life, survival and breeding investment (clutch size), indicated that urban birds generally have higher survival, but smaller clutch sizes. The latter finding (smaller clutches in urban habitats) seemed to be mainly a characteristic of smaller passerines. We also reviewed urbanization studies on other traits that can be associated with pace-of-life and are related to either reproductive investment or self-maintenance. Though sample sizes were generally too small to conduct formal meta-analyses, published literature suggests that urban birds tend to produce lower-quality sexual signals and invest more in offspring care. The latter finding is in agreement with the adult survival hypothesis, proposing that higher adult survival prospects favour investment in fewer offspring per year. According to our hypothesis, differences in age structure should arise between urban and rural populations, providing a novel alternative explanation for physiological differences and earlier breeding. We encourage more research investigating how telomere dynamics, immune defences, antioxidants and oxidative damage in different tissues vary along the urbanization gradient, and suggest that applying pace-of-life framework to studies of variation in physiological traits along the urbanization gradient might be the next direction to improve our understanding of urbanization as an evolutionary process.