Identifying potential evolutionary consequences of climate-driven phenological shifts

Climate change is shifting the phenology of many species throughout the world. While the interspecific consequences of these phenological shifts have been well documented, the intraspecific shifts and their resultant evolutionary consequences remain relatively unexplored. Here, we present a conceptual framework and overview of how phenological shifts within species can drive evolutionary change. We suggest that because the impacts of climate change are likely to vary across the range of a species and differentially impact individuals, phenological shifts may often be highly variable both within and among populations. Together these changes have the potential to alter existing patterns of gene flow and influence evolutionary trajectories by increasing phenological isolation and connectivity. Recent research examining the response of species to contemporary climate change suggests that both phenological isolation and connectivity may be likely responses to future climate change. However, recent studies also show mixed results on whether adaptive responses to climate change are likely to occur, as some populations have already shown adaptive responses to changing climate, while others have not despite fitness costs. While predicting the exact consequences of intraspecific phenological shifts may be difficult, identifying the evolutionary implications of these shifts will allow a better understanding of the effects of future climate change on species persistence and adaptation.     

Biological responses to environmental heterogeneity under future ocean conditions

Organisms are projected to face unprecedented rates of change in future ocean conditions due to anthropogenic climate‐change. At present, marine life encounters a wide range of environmental heterogeneity from natural fluctuations to mean climate change. Manipulation studies suggest that biota from more variable marine environments have more phenotypic plasticity to tolerate environmental heterogeneity. Here, we consider current strategies employed by a range of representative organisms across various habitats – from short‐lived phytoplankton to long‐lived corals – in response to environmental heterogeneity. We then discuss how, if and when organismal responses (acclimate/migrate/adapt) may be altered by shifts in the magnitude of the mean climate‐change signal relative to that for natural fluctuations projected for coming decades. The findings from both novel climate‐change modelling simulations and prior biological manipulation studies, in which natural fluctuations are superimposed on those of mean change, provide valuable insights into organismal responses to environmental heterogeneity. Manipulations reveal that different experimental outcomes are evident between climate‐change treatments which include natural fluctuations vs. those which do not. Modelling simulations project that the magnitude of climate variability, along with mean climate change, will increase in coming decades, and hence environmental heterogeneity will increase, illustrating the need for more realistic biological manipulation experiments that include natural fluctuations. However, simulations also strongly suggest that the timescales over which the mean climate‐change signature will become dominant, relative to natural fluctuations, will vary for individual properties, being most rapid for CO₂ (~10 years from present day) to 4 decades for nutrients. We conclude that the strategies used by biota to respond to shifts in environmental heterogeneity may be complex, as they will have to physiologically straddle wide‐ranging timescales in the alteration of ocean conditions, including the need to adapt to rapidly rising CO₂ and also acclimate to environmental heterogeneity in more slowly changing properties such as warming.     

Effects of past and projected river discharge variability on freshwater production in an anadromous fish

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Influence of hydrologic attributes on brown trout recruitment in low-latitude range margins

Factors controlling brown trout Salmo trutta recruitment in Mediterranean areas are largely unknown, despite the relevance this may have for fisheries management. The effect of hydrological variability on survival of young brown trout was studied during seven consecutive years in five resident populations from the southern range of the species distribution. Recruit density at the end of summer varied markedly among year-classes and rivers during the study period. Previous work showed that egg density the previous fall did not account for more than 50% of the observed variation in recruitment density. Thus, we expected that climatic patterns, as determinants of discharge and water temperature, would play a role in the control of young trout abundance. We tested this by analyzing the effects of flow variation and predictability on young trout survival during the spawning to emergence and the summer drought periods. Both hatching and emergence times and length of hatching and emergence periods were similar between years within each river but varied considerably among populations, due to differences in water temperature. Interannual variation in flow attributes during spawning to emergence and summer drought affected juvenile survival in all populations, once the effect of endogenous factors was removed. Survival rate was significantly related to the timing, magnitude and duration of extreme water conditions, and to the rate of change in discharge during hatching and emergence times in most rivers. The magnitude and duration of low flows during summer drought appeared to be a critical factor for survival of young trout. Our findings suggest that density-independent factors, i.e., hydrological variability, play a central role in the population dynamics of brown trout in populations from low-latitude range margins. Reported effects of hydrologic attributes on trout survival are likely to be increasingly important if, as predicted, climate change leads to greater extremes and variability of flow regimes.     

Have bird distributions shifted along an elevational gradient on a tropical mountain?

An upward shift in elevation is one of the most conspicuous species responses to climate change. Nevertheless, downward shifts and, apparently, the absences of response have also been recently reported. Given the growing evidence of multiple responses of species distributions due to climate change and the paucity of studies in the tropics, we evaluated the response of a montane bird community to climate change, without the confounding effects of land‐use change. To test for elevational shifts, we compared the distribution of 21 avian species in 1998 and 2015 using occupancy models. The historical data set was based on point counts, whereas the contemporary data set was based on acoustic monitoring. We detected a similar number of species in historical (36) and contemporary data sets (33). We show an overall pattern of no significant change in range limits for most species, although there was a significant shift in the range limit of eight species (38%). Elevation limits shifted mostly upward, and this pattern was more common for upper than lower limits. Our results highlight the variability of species responses to climate change and illustrate how acoustic monitoring provides an easy and powerful way to monitor animal populations along elevational gradients.     

Spatial and temporal heterogeneity in climate change limits species' dispersal capabilities and adaptive potential

Global climate change has already caused local declines and extinctions. These losses are generally thought to occur because climate change is progressing too rapidly for populations to keep pace. Based on this hypothesis, numerous predictive frameworks have been developed to project future range shifts and changes in population dynamics resulting from global climate change. However, recent empirical work has demonstrated that seasonally asynchronous climate change regimes – when a region is warming during some parts of the year, but cooling in others – are constraining species' responses to climate change more strongly than rapid warming, leading to intra‐specific variation in responses to climate change and local population declines. Here, we couple a review of the literature related to asynchronous climate change regimes with meta‐population simulations and an analysis of long‐term North American climate trends to show that seasonally asynchronous regimes are occurring throughout most of North America and that their current spatial distribution may be a strong barrier to dispersal and gene flow across many species' ranges. Thus, even though adaptation to climate change may potentially be more common and rapid than previously thought, species whose ranges overlap with asynchronous regimes will likely succumb to local declines that may be difficult to mitigate via dispersal. Future climate‐related predictive frameworks should therefore incorporate asynchronous regimes as well as more traditional measures of climate velocity in order to fully capture the array of potential future climate change scenarios.     

森林干扰对大兴安岭北部森林小流域径流情势的影响

为了解大兴安岭森林流域水文过程对森林干扰的响应,利用近配对流域方法,排除了气候变量的时空差异,对比研究了森林干扰后大兴安岭北部典型森林小流域(100 km2)洪峰径流(High flow)和枯水径流(Low flow)径流情势(Flow regimes)的变化趋势。结果表明,森林干扰对枯水径流情势影响显著,与对照流域(小北沟流域)相比,森林干扰(占流域总面积的6.74%)使老沟河流域平均枯水径流流量降低了26.58%,平均枯水径流变异系数值增加了36.77%,并且差异达到极显著水平(P0.01)。另一方面,森林植被的干扰相对增加了森林小流域的洪峰流量、历时和变异性,但与对照流域相比差异均未达到统计显著水平,说明小面积的森林植被干扰未能引起流域洪峰径流情势的显著变化。进一步对配对流域的径流浮动系数(Flashiness Index)的分析发现,森林干扰显著增加了森林小流域的径流浮动性,研究时段内干扰流域的径流浮动系数为0.078,是对照流域(0.057)的1.37倍。大兴安岭北部森林小流域的天然径流情势(Natural flow regimes)对森林干扰比较敏感,在与水文循环联系紧密的区域(例如河岸带),小范围的森林干扰便可以引起径流情势的显著变化,这在未来该地区森林和水资源的管理中需要特别注意。     

Environmental variability and population dynamics: do European and North American ducks play by the same rules?

Density dependence, population regulation, and variability in population size are fundamental population processes, the manifestation and interrelationships of which are affected by environmental variability. However, there are surprisingly few empirical studies that distinguish the effect of environmental variability from the effects of population processes. We took advantage of a unique system, in which populations of the same duck species or close ecological counterparts live in highly variable (north American prairies) and in stable (north European lakes) environments, to distinguish the relative contributions of environmental variability (measured as between‐year fluctuations in wetland numbers) and intraspecific interactions (density dependence) in driving population dynamics. We tested whether populations living in stable environments (in northern Europe) were more strongly governed by density dependence than populations living in variable environments (in North America). We also addressed whether relative population dynamical responses to environmental variability versus density corresponded to differences in life history strategies between dabbling (relatively “fast species” and governed by environmental variability) and diving (relatively “slow species” and governed by density) ducks. As expected, the variance component of population fluctuations caused by changes in breeding environments was greater in North America than in Europe. Contrary to expectations, however, populations in more stable environments were not less variable nor clearly more strongly density dependent than populations in highly variable environments. Also, contrary to expectations, populations of diving ducks were neither more stable nor stronger density dependent than populations of dabbling ducks, and the effect of environmental variability on population dynamics was greater in diving than in dabbling ducks. In general, irrespective of continent and species life history, environmental variability contributed more to variation in species abundances than did density. Our findings underscore the need for more studies on populations of the same species in different environments to verify the generality of current explanations about population dynamics and its association with species life history.     

Beyond average: an experimental test of temperature variability on the population dynamics of <Emphasis Type="Italic">Tribolium confusum</Emphasis>

The relationship between ectotherm ecology and climatic conditions has been mainly evaluated in terms of average conditions. Average temperature is the more common climatic variable used in physiological and population studies, and its effect on individual and population-level processes is well understood. However, the intrinsic variability of thermal conditions calls attention to the potential effects that this variability could have in ecological systems. Regarding this point, two hypotheses are proposed. From the allocation principle, it may be inferred that if temperature variability is high enough to induce stress in the organisms, then this extra-cost should reduce the energetic budget for reproduction, which will be reflected in population parameters. Moreover, a mathematical property of non-linear functions, Jensen’s inequality, indicates that, in concave functions, like the temperature–reproduction performance function, variability reduces the expected value of the output variable, and again modifies population parameters. To test these hypotheses, experimental cultures of Tribolium confusum under two different thermal variability regimens were carried out. With these data, we fitted a simple population dynamics model to evaluate the predictions of our hypothesis. The results show that thermal variability reduces the maximum reproductive rate of the population but no other parameters such as carrying capacity or the nonlinear factor in a nonlinear version of the Ricker model, which confirms our hypotheses. This result has important consequences, such as the paradoxical increase in population variability under a decrease in thermal variability and the necessary incorporation of climatic variability to evaluate the net effect of climate change on the dynamics of natural populations.     

Local Variability Mediates Vulnerability of Trout Populations to Land Use and Climate Change

Land use and climate change occur simultaneously around the globe. Fully understanding their separate and combined effects requires a mechanistic understanding at the local scale where their effects are ultimately realized. Here we applied an individual-based model of fish population dynamics to evaluate the role of local stream variability in modifying responses of Coastal Cutthroat Trout (Oncorhynchus clarkii clarkii) to scenarios simulating identical changes in temperature and stream flows linked to forest harvest, climate change, and their combined effects over six decades. We parameterized the model for four neighboring streams located in a forested headwater catchment in northwestern Oregon, USA with multi-year, daily measurements of stream temperature, flow, and turbidity (2007–2011), and field measurements of both instream habitat structure and three years of annual trout population estimates. Model simulations revealed that variability in habitat conditions among streams (depth, available habitat) mediated the effects of forest harvest and climate change. Net effects for most simulated trout responses were different from or less than the sum of their separate scenarios. In some cases, forest harvest countered the effects of climate change through increased summer flow. Climate change most strongly influenced trout (earlier fry emergence, reductions in biomass of older trout, increased biomass of young-of-year), but these changes did not consistently translate into reductions in biomass over time. Forest harvest, in contrast, produced fewer and less consistent responses in trout. Earlier fry emergence driven by climate change was the most consistent simulated response, whereas survival, growth, and biomass were inconsistent. Overall our findings indicate a host of local processes can strongly influence how populations respond to broad scale effects of land use and climate change.     

Where will species go? Incorporating new advances in climate modelling into projections of species distributions

Bioclimatic models are the primary tools for simulating the impact of climate change on species distributions. Part of the uncertainty in the output of these models results from uncertainty in projections of future climates. To account for this, studies often simulate species responses to climates predicted by more than one climate model and/or emission scenario. One area of uncertainty, however, has remained unexplored: internal climate model variability. By running a single climate model multiple times, but each time perturbing the initial state of the model slightly, different but equally valid realizations of climate will be produced. In this paper, we identify how ongoing improvements in climate models can be used to provide guidance for impacts studies. In doing so we provide the first assessment of the extent to which this internal climate model variability generates uncertainty in projections of future species distributions, compared with variability between climate models. We obtained data on 13 realizations from three climate models (three from CSIRO Mark2 v3.0, four from GISS AOM, and six from MIROC v3.2) for two time periods: current (1985–1995) and future (2025–2035). Initially, we compared the simulated values for each climate variable (P, T_max, T_min, and T_mean) for the current period to observed climate data. This showed that climates simulated by realizations from the same climate model were more similar to each other than to realizations from other models. However, when projected into the future, these realizations followed different trajectories and the values of climate variables differed considerably within and among climate models. These had pronounced effects on the projected distributions of nine Australian butterfly species when modelled using the BIOCLIM component of DIVA-GIS. Our results show that internal climate model variability can lead to substantial differences in the extent to which the future distributions of species are projected to change. These can be greater than differences resulting from between-climate model variability. Further, different conclusions regarding the vulnerability of species to climate change can be reached due to internal model variability. Clearly, several climate models, each represented by multiple realizations, are required if we are to adequately capture the range of uncertainty associated with projecting species distributions in the future.     

Clinal adaptation and adaptive plasticity in Artemisia californica: implications for the response of a foundation species to predicted climate change

Local adaptation and plasticity pose significant obstacles to predicting plant responses to future climates. Although local adaptation and plasticity in plant functional traits have been documented for many species, less is known about population‐level variation in plasticity and whether such variation is driven by adaptation to environmental variation. We examined clinal variation in traits and performance – and plastic responses to environmental change – for the shrub Artemisia californica along a 700 km gradient characterized (from south to north) by a fourfold increase in precipitation and a 61% decrease in interannual precipitation variation. Plants cloned from five populations along this gradient were grown for 3 years in treatments approximating the precipitation regimes of the north and south range margins. Most traits varying among populations did so clinally; northern populations (vs. southern) had higher water‐use efficiencies and lower growth rates, C : N ratios and terpene concentrations. Notably, there was variation in plasticity for plant performance that was strongly correlated with source site interannual precipitation variability. The high‐precipitation treatment (vs. low) increased growth and flower production more for plants from southern populations (181% and 279%, respectively) than northern populations (47% and 20%, respectively). Overall, precipitation variability at population source sites predicted 86% and 99% of variation in plasticity in growth and flowering, respectively. These striking, clinal patterns in plant traits and plasticity are indicative of adaptation to both the mean and variability of environmental conditions. Furthermore, our analysis of long‐term coastal climate data in turn indicates an increase in interannual precipitation variation consistent with most global change models and, unexpectedly, this increased variation is especially pronounced at historically stable, northern sites. Our findings demonstrate the critical need to integrate fundamental evolutionary processes into global change models, as contemporary patterns of adaptation to environmental clines will mediate future plant responses to projected climate change.     

Analysis of climate paths reveals potential limitations on species range shifts

Forecasts of species endangerment under climate change usually ignore the processes by which species ranges shift. By analysing the 'climate paths' that range shifts might follow, and two key range-shift processes--dispersal and population persistence--we show that short-term climatic and population characteristics have dramatic effects on range-shift forecasts. By employing this approach with 15 amphibian species in the western USA, we make unexpected predictions. First, inter-decadal variability in climate change can prevent range shifts by causing gaps in climate paths, even in the absence of geographic barriers. Second, the hitherto unappreciated trait of persistence during unfavourable climatic conditions is critical to species range shifts. Third, climatic fluctuations and low persistence could lead to endangerment even if the future potential range size is large. These considerations may render habitat corridors ineffectual for some species, and conservationists may need to consider managed relocation and augmentation of in situ populations.     

Patterns of drought tolerance in major European temperate forest trees: climatic drivers and levels of variability

The future performance of native tree species under climate change conditions is frequently discussed, since increasingly severe and more frequent drought events are expected to become a major risk for forest ecosystems. To improve our understanding of the drought tolerance of the three common European temperate forest tree species Norway spruce, silver fir and common beech, we tested the influence of climate and tree‐specific traits on the inter and intrasite variability in drought responses of these species. Basal area increment data from a large tree‐ring network in Southern Germany and Alpine Austria along a climatic cline from warm‐dry to cool‐wet conditions were used to calculate indices of tolerance to drought events and their variability at the level of individual trees and populations. General patterns of tolerance indicated a high vulnerability of Norway spruce in comparison to fir and beech and a strong influence of bioclimatic conditions on drought response for all species. On the level of individual trees, low‐growth rates prior to drought events, high competitive status and low age favored resilience in growth response to drought. Consequently, drought events led to heterogeneous and variable response patterns in forests stands. These findings may support the idea of deliberately using spontaneous selection and adaption effects as a passive strategy of forest management under climate change conditions, especially a strong directional selection for more tolerant individuals when frequency and intensity of summer droughts will increase in the course of global climate change.     

Desert grassland responses to climate and soil moisture suggest divergent vulnerabilities across the southwestern United States

Climate change predictions include warming and drying trends, which are expected to be particularly pronounced in the southwestern United States. In this region, grassland dynamics are tightly linked to available moisture, yet it has proven difficult to resolve what aspects of climate drive vegetation change. In part, this is because it is unclear how heterogeneity in soils affects plant responses to climate. Here, we combine climate and soil properties with a mechanistic soil water model to explain temporal fluctuations in perennial grass cover, quantify where and the degree to which incorporating soil water dynamics enhances our ability to understand temporal patterns, and explore the potential consequences of climate change by assessing future trajectories of important climate and soil water variables. Our analyses focused on long‐term (20–56 years) perennial grass dynamics across the Colorado Plateau, Sonoran, and Chihuahuan Desert regions. Our results suggest that climate variability has negative effects on grass cover, and that precipitation subsidies that extend growing seasons are beneficial. Soil water metrics, including the number of dry days and availability of water from deeper (>30 cm) soil layers, explained additional grass cover variability. While individual climate variables were ranked as more important in explaining grass cover, collectively soil water accounted for 40–60% of the total explained variance. Soil water conditions were more useful for understanding the responses of C₃ than C₄ grass species. Projections of water balance variables under climate change indicate that conditions that currently support perennial grasses will be less common in the future, and these altered conditions will be more pronounced in the Chihuahuan Desert and Colorado Plateau. We conclude that incorporating multiple aspects of climate and accounting for soil variability can improve our ability to understand patterns, identify areas of vulnerability, and predict the future of desert grasslands.     

Climate variability and vulnerability to climate change: a review

The focus of the great majority of climate change impact studies is on changes in mean climate. In terms of climate model output, these changes are more robust than changes in climate variability. By concentrating on changes in climate means, the full impacts of climate change on biological and human systems are probably being seriously underestimated. Here, we briefly review the possible impacts of changes in climate variability and the frequency of extreme events on biological and food systems, with a focus on the developing world. We present new analysis that tentatively links increases in climate variability with increasing food insecurity in the future. We consider the ways in which people deal with climate variability and extremes and how they may adapt in the future. Key knowledge and data gaps are highlighted. These include the timing and interactions of different climatic stresses on plant growth and development, particularly at higher temperatures, and the impacts on crops, livestock and farming systems of changes in climate variability and extreme events on pest‐weed‐disease complexes. We highlight the need to reframe research questions in such a way that they can provide decision makers throughout the food system with actionable answers, and the need for investment in climate and environmental monitoring. Improved understanding of the full range of impacts of climate change on biological and food systems is a critical step in being able to address effectively the effects of climate variability and extreme events on human vulnerability and food security, particularly in agriculturally based developing countries facing the challenge of having to feed rapidly growing populations in the coming decades.     

Morphological plasticity in a native freshwater fish from semiarid Australia in response to variable water flows

In fishes, alterations to the natural flow regime are associated with divergence in body shape morphology compared with individuals from unaltered habitats. However, it is unclear whether this morphological divergence is attributable to evolutionary responses to modified flows, or is a result of phenotypic plasticity. Fishes inhabiting arid regions are ideal candidates for studying morphological plasticity as they are frequently exposed to extreme natural hydrological variability. We examined the effect of early exposure to flows on the development of body shape morphology in the western rainbowfish (Melanotaenia australis), a freshwater fish that is native to semiarid northwest Australia. Wild fish were collected from a region (the Hamersley Ranges) where fish in some habitats are subject to altered water flows due to mining activity. The offspring of wild‐caught fish were reared in replicated fast‐flow or slow‐flow channels, and geometric morphometric analyses were used to evaluate variation in fish body shape following 3, 6, 9, and 12 months of exposure. Water flows influenced fish morphology after 6 and 9 months of flow exposure, with fish in fast‐flow environments displaying a more robust body shape than those in slow‐flow habitats. No effect of flow exposure was observed at 3 and 12 months. Fishes also showed significant morphological variation within flow treatments, perhaps due to subtle differences in water flow among the replicate channels. Our findings suggest that early exposure to water flows can induce shifts in body shape morphology in arid zone freshwater fishes. Morphological plasticity may act to buffer arid zone populations from the impacts of anthropogenic activities, but further studies are required to link body shape plasticity with behavioral performance in habitats with modified flows.     

Directionality of recent bird distribution shifts and climate change in Great Britain

There is good evidence that species' distributions are shifting poleward in response to climate change and wide interest in the magnitude of such responses for scientific and conservation purposes. It has been suggested from the directions of climatic changes that species' distribution shifts may not be simply poleward, but this has been rarely tested with observed data. Here, we apply a novel approach to measuring range shifts on axes ranging through 360°, to recent data on the distributions of 122 species of British breeding birds during 1988–1991 and 2008–2011. Although previously documented poleward range shifts have continued, with an average 13.5 km shift northward, our analysis indicates this is an underestimate because it ignores common and larger shifts that occurred along axes oriented to the north‐west and north‐east. Trailing edges contracted from a broad range of southerly directions. Importantly, these results are derived from systematically collected data so confounding observer‐effort biases can be discounted. Analyses of climate for the same period show that whilst temperature trends should drive species along a north–north‐westerly trajectory, directional responses to precipitation will depend on both the time of year that is important for determining a species' distribution, and the location of the range margin. Directions of species' range centroid shift were not correlated with spatial trends in any single climate variable. We conclude that range shifts of British birds are multidirectional, individualistic and probably determined by species‐specific interactions of multiple climate factors. Climate change is predicted to lead to changes in community composition through variation in the rates that species' ranges shift; our results suggest communities could change further owing to constituent species shifting along different trajectories. We recommend more studies consider directionality in climate and range dynamics to produce more appropriate measures of observed and expected responses to climate change.     

Terrestrial insects and climate change: adaptive responses in key traits

Understanding and predicting how adaptation will contribute to species' resilience to climate change will be paramount to successfully managing biodiversity for conservation, agriculture, and human health‐related purposes. Making predictions that capture how species will respond to climate change requires an understanding of how key traits and environmental drivers interact to shape fitness in a changing world. Current trait‐based models suggest that low‐ to mid‐latitude populations will be most at risk, although these models focus on upper thermal limits, which may not be the most important trait driving species' distributions and fitness under climate change. In this review, we discuss how different traits (stress, fitness and phenology) might contribute and interact to shape insect responses to climate change. We examine the potential for adaptive genetic and plastic responses in these key traits and show that, although there is evidence of range shifts and trait changes, explicit consideration of what underpins these changes, be that genetic or plastic responses, is largely missing. Despite little empirical evidence for adaptive shifts, incorporating adaptation into models of climate change resilience is essential for predicting how species will respond under climate change. We are making some headway, although more data are needed, especially from taxonomic groups outside of Drosophila, and across diverse geographical regions. Climate change responses are likely to be complex, and such complexity will be difficult to capture in laboratory experiments. Moving towards well designed field experiments would allow us to not only capture this complexity, but also study more diverse species.