100 million years of turtle paleoniche dynamics enable the prediction of latitudinal range shifts in a warming world

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Climate change, anthropogenic disturbance and the northward range expansion of Lactuca serriola (Asteraceae)

Aim The distribution range of Lactuca serriola, a species native to the summer‐dry mediterranean climate, has expanded northwards during the last 250 years. This paper assesses the influence of climate on the range expansion of this species and highlights the importance of anthropogenic disturbance to its spread. Location Central and Northern Europe. Methods Data on the geographic distribution of L. serriola were assembled through a literature search as well as through floristic and herbarium surveys. Maps of the spread of L. serriola in Central and Northern Europe were prepared based on herbarium data. The spread was assessed more precisely in Germany, Austria and Great Britain by pooling herbarium and literature data. We modelled the bioclimatic niche of the species using occurrence and climatic data covering the last century to generate projections of suitable habitats under the climatic conditions of five time periods. We tested whether the observed distribution of L. serriola could be explained for each time period, assuming that the climatic niche of the species was conserved across time. Results The species has spread northwards since the beginning of the 19th century. We show that climate warming in Europe increased the number of sites suitable for the species at northern latitudes. Until the late 1970s, the distribution of the species corresponded to the climatically suitable sites available. For the last two decades, however, we could not show any significant relationship between the increase in suitable sites and the distributional range change of L. serriola. However, we highlight potential areas the species could spread to in the future (Great Britain, southern Scandinavia and the Swedish coast). It is predominantly non‐climatic influences of global change that have contributed to its rapid spread. Main conclusions The observation that colonizing species are not filling their climatically suitable range might imply that, potentially, other ruderal species could expand far beyond their current range. Our work highlights the importance of historical floristic and herbarium data for understanding the expansion of a species. Such historical distributional data can provide valuable information for those planning the management of contemporary environmental problems, such as species responses to environmental change.     

Climate change and fire effects on a prairie–woodland ecotone: projecting species range shifts with a dynamic global vegetation model

Large shifts in species ranges have been predicted under future climate scenarios based primarily on niche‐based species distribution models. However, the mechanisms that would cause such shifts are uncertain. Natural and anthropogenic fires have shaped the distributions of many plant species, but their effects have seldom been included in future projections of species ranges. Here, we examine how the combination of climate and fire influence historical and future distributions of the ponderosa pine–prairie ecotone at the edge of the Black Hills in South Dakota, USA, as simulated by MC1, a dynamic global vegetation model that includes the effects of fire, climate, and atmospheric CO₂ concentration on vegetation dynamics. For this purpose, we parameterized MC1 for ponderosa pine in the Black Hills, designating the revised model as MC1‐WCNP. Results show that fire frequency, as affected by humidity and temperature, is central to the simulation of historical prairies in the warmer lowlands versus woodlands in the cooler, moister highlands. Based on three downscaled general circulation model climate projections for the 21st century, we simulate greater frequencies of natural fire throughout the area due to substantial warming and, for two of the climate projections, lower relative humidity. However, established ponderosa pine forests are relatively fire resistant, and areas that were initially wooded remained so over the 21st century for most of our future climate x fire management scenarios. This result contrasts with projections for ponderosa pine based on climatic niches, which suggest that its suitable habitat in the Black Hills will be greatly diminished by the middle of the 21st century. We hypothesize that the differences between the future predictions from these two approaches are due in part to the inclusion of fire effects in MC1, and we highlight the importance of accounting for fire as managed by humans in assessing both historical species distributions and future climate change effects.     

US forest response to projected climate‐related stress: a tolerance perspective

Although it is widely recognized that climate change will require a major spatial reorganization of forests, our ability to predict exactly how and where forest characteristics and distributions will change has been rather limited. Current efforts to predict future distribution of forested ecosystems as a function of climate include species distribution models (for fine‐scale predictions) and potential vegetation climate envelope models (for coarse‐grained, large‐scale predictions). Here, we develop and apply an intermediate approach wherein we use stand‐level tolerances of environmental stressors to understand forest distributions and vulnerabilities to anticipated climate change. In contrast to other existing models, this approach can be applied at a continental scale while maintaining a direct link to ecologically relevant, climate‐related stressors. We first demonstrate that shade, drought, and waterlogging tolerances of forest stands are strongly correlated with climate and edaphic conditions in the conterminous United States. This discovery allows the development of a tolerance distribution model (TDM), a novel quantitative tool to assess landscape level impacts of climate change. We then focus on evaluating the implications of the drought TDM. Using an ensemble of 17 climate change models to drive this TDM, we estimate that 18% of US ecosystems are vulnerable to drought‐related stress over the coming century. Vulnerable areas include mostly the Midwest United States and Northeast United States, as well as high‐elevation areas of the Rocky Mountains. We also infer stress incurred by shifting climate should create an opening for the establishment of forest types not currently seen in the conterminous United States.     

Soil drying and nitrogen availability modulate carbon and water exchange over a range of annual precipitation totals and grassland vegetation types

Increased intensity in precipitation events and longer periods of water deficit are predicted as a general trend under future climate scenarios with potentially large effects on terrestrial ecosystem function. The primary objective of this study was to understand how variation in the intensity of precipitation inputs followed by intermittent soil drying events influence leaf and ecosystem carbon dioxide (CO₂) and water exchange in a California annual grassland mesocosm experiment. We further examined how nitrogen (N) availability, and differences in plant community composition (grass-forb combinations) affected gas exchange responses to the precipitation treatments. Net ecosystem CO₂ exchange (NEE) and evapotranspiration (ET) increased significantly with greater precipitation and were positively correlated with soil moisture. A repeated 10-day soil drying period, following 11 days of watering, strongly depressed NEE over a range of annual precipitation totals (297, 657 and 987 mm), and plant community types. Ecosystem dark respiration ( R _e) and leaf level photosynthesis ( A _max) showed greater sensitivity to periods of soil drying in the low precipitation plots (297 mm). N additions significantly increased NEE and R _e, particularly as water availability was increased. Across the range of precipitation totals and plant community types, intermittent periods of soil moisture deficit and native soil N availability constrained leaf and ecosystem level CO₂ exchange, while the influence on water vapor exchange was less pronounced.     

Climatic variation and simulated patterns in seedling establishment of two dominant grasses at a semi‐arid‐arid grassland ecotone

Abstract. The objective of this study was to predict the effects of climatic variation at multiple temporal frequencies on seedling establishment by two congeneric C₄ perennial grasses (Bouteloua gracilis and B. eriopoda) at the ecotone between shortgrass steppe grassland and Chihuahuan desert grassland in central New Mexico, USA. The approach was to use a daily time‐step simulation model to determine the occurrence of a recruitment event in each year based upon the amount and timing of soil water required for establishment. Historical weather data were used to predict effects of seasonal and inter‐decadal variation in climate on establishment. A sensitivity analysis was used to predict effects of directional climate change on establishment. Bouteloua gracilis had a broad pattern of simulated establishment from May through September that included periods with high year‐to‐year variation in precipitation. B. eriopoda establishment events occurred primarily in July when precipitation amounts were most reliable. Climatic conditions from 1949 through 1968 were more favorable for B. eriopoda establishment compared to the cooler, wetter conditions from 1969 through 1988 that favored B. gracilis. Establishment of B. eriopoda was lowest in El Niño years whereas B. gracilis establishment was highest in La Niña years. Establishment of B. gracilis was most sensitive to temperature when precipitation was higher than current amounts. The greatest response to temperature by B. eriopoda for all precipitation amounts occurred at cooler temperatures than found currently. These results indicate that climatic variation at multiple frequencies has differential effects on seedling establishment for these two perennial grasses, and may account at least in part for patterns in dominance at this biome transition zone.     

Insect distribution in response to climate change based on a model: Review of function and use of CLIMEX

The significant dependence of agricultural productivity on pest control requires pest distribution predictions at an early stage of pest invasion. Because pest cycles are critically affected by climate, climate is one of the most important factors for predicting an invasive pest. CLIMEX is a highly effective tool that can predict potential geographical species distributions, and test the regional suitability for a target species' habitat based on data including climate change scenarios. CLIMEX has been recently used in Europe, North America, China and Australia, among others. However, for modeling species distributions in Korea, the use of the model has been limited to date. This study aimed to first introduce the function and application of CLIMEX by reviewing important studies using this model. Second, we investigated previous studies using the model simulation to demonstrate the practical applicability of CLIMEX for the agricultural sector, and its use in forecasting.     

Leaf hydraulic vulnerability to drought is linked to site water availability across a broad range of species and climates

Background and Aims

Vulnerability of the leaf hydraulic pathway to water-stress-induced dysfunction is a key component of drought tolerance in plants and may be important in defining species'' climatic range. However, the generality of the association between leaf hydraulic vulnerability and climate across species and sites remains to be tested.

Methods

Leaf hydraulic vulnerability to drought (P50_leaf, the water potential inducing 50 % loss in hydraulic function) was measured in a diverse group of 92 woody, mostly evergreen angiosperms from sites across a wide range of habitats. These new data together with some previously published were tested against key climate indices related to water availability. Differences in within-site variability in P50_leaf between sites were also examined.

Key Results

Values of hydraulic vulnerability to drought in leaves decreased strongly (i.e. became more negative) with decreasing annual rainfall and increasing aridity across sites. The standard deviation in P50_leaf values recorded within each site was positively correlated with increasing aridity. P50_leaf was also a good indicator of the climatic envelope across each species'' distributional range as well as their dry-end distributional limits within Australia, although this relationship was not consistently detectable within sites.

Conclusions

The findings indicate that species sorting processes have influenced distributional patterns of P50_leaf across the rainfall spectrum, but alternative strategies for dealing with water deficit exist within sites. The strong link to aridity suggests leaf hydraulic vulnerability may influence plant distributions under future climates.     

Historical and event‐based bioclimatic suitability predicts regional forest vulnerability to compound effects of severe drought and bark beetle infestation

Vulnerability to climate change, and particularly to climate extreme events, is expected to vary across species ranges. Thus, we need tools to standardize the variability in regional climatic legacy and extreme climate across populations and species. Extreme climate events (e.g., droughts) can erode populations close to the limits of species' climatic tolerance. Populations in climatic‐core locations may also become vulnerable because they have developed a greater demand for resources (i.e., water) that cannot be enough satisfied during the periods of scarcity. These mechanisms can become exacerbated in tree populations when combined with antagonistic biotic interactions, such as insect infestation. We used climatic suitability indices derived from Species Distribution Models (SDMs) to standardize the climatic conditions experienced across Pinus edulis populations in southwestern North America, during a historical period (1972–2000) and during an extreme event (2001–2007), when the compound effect of hot drought and bark beetle infestation caused widespread die‐off and mortality. Pinus edulis climatic suitability diminished dramatically during the die‐off period, with remarkable variation between years. P. edulis die‐off occurred mainly not just in sites that experienced lower climatic suitability during the drought but also where climatic suitability was higher during the historical period. The combined effect of historically high climatic suitability and a marked decrease in the climatic suitability during the drought best explained the range‐wide mortality. Lagged effects of climatic suitability loss in previous years and co‐occurrence of Juniperus monosperma also explained P. edulis die‐off in particular years. Overall, the study shows that past climatic legacy, likely determining acclimation, together with competitive interactions plays a major role in responses to extreme drought. It also provides a new approach to standardize the magnitude of climatic variability across populations using SDMs, improving our capacity to predict population's or species' vulnerability to climatic change.     

Demographic consequences of climate change and land cover help explain a history of extirpations and range contraction in a declining snake species

Developing conservation strategies for threatened species increasingly requires understanding vulnerabilities to climate change, in terms of both demographic sensitivities to climatic and other environmental factors, and exposure to variability in those factors over time and space. We conducted a range‐wide, spatially explicit climate change vulnerability assessment for Eastern Massasauga (Sistrurus catenatus), a declining endemic species in a region showing strong environmental change. Using active season and winter adult survival estimates derived from 17 data sets throughout the species' range, we identified demographic sensitivities to winter drought, maximum precipitation during the summer, and the proportion of the surrounding landscape dominated by agricultural and urban land cover. Each of these factors was negatively associated with active season adult survival rates in binomial generalized linear models. We then used these relationships to back‐cast adult survival with dynamic climate variables from 1950 to 2008 using spatially explicit demographic models. Demographic models for 189 population locations predicted known extant and extirpated populations well (AUC = 0.75), and models based on climate and land cover variables were superior to models incorporating either of those effects independently. These results suggest that increasing frequencies and severities of extreme events, including drought and flooding, have been important drivers of the long‐term spatiotemporal variation in a demographic rate. We provide evidence that this variation reflects nonadaptive sensitivity to climatic stressors, which are contributing to long‐term demographic decline and range contraction for a species of high‐conservation concern. Range‐wide demographic modeling facilitated an understanding of spatial shifts in climatic suitability and exposure, allowing the identification of important climate refugia for a dispersal‐limited species. Climate change vulnerability assessment provides a framework for linking demographic and distributional dynamics to environmental change, and can thereby provide unique information for conservation planning and management.     

Testing the limits of resistance: a 19‐year study of Mediterranean grassland response to grazing regimes

A synthesis of a long‐term (19 years) study assessing the effects of cattle grazing on the structure and composition of a Mediterranean grassland in north‐eastern Israel is presented, with new insights into the response of the vegetation to grazing management and rainfall. We hypothesized that the plant community studied would be resistant to high grazing intensities and rainfall variability considering the combined long history of land‐use and unpredictable climatic conditions where this community evolved. Treatments included manipulations of stocking densities (moderate, heavy, and very heavy) and of grazing regimes (continuous vs. seasonal), in a factorial design. The effect of interannual rainfall variation on the expression of grazing impacts on the plant community was minor. The main effects of grazing on relative cover of plant functional groups were related to early vs. late seasonal grazing. Species diversity and equitability were remarkably stable across all grazing treatments. A reduction in tall grass cover at higher stocking densities was correlated with increased cover of less palatable groups such as annual and perennial thistles, as well as shorter and prostrate groups such as short annual grasses. This long‐term study shows that interannual fluctuations in plant functional group composition could be partly accounted for by grazing pressure and timing, but not by the measured rainfall variables. Grazing affected the dominance of tall annual grasses. However, the persistence of tall grasses and more palatable species over time, despite large differences in grazing pressure and timing, supports the idea that Mediterranean grasslands are highly resistant to prolonged grazing. Indeed, even under the most extreme grazing conditions applied, there were no signs of deterioration or collapse of the ecosystem. This high resistance to grazing intensity and interannual fluctuation in climatic conditions should favor the persistence of the plant community under forecasted increasing unpredictability due to climate change.     

Annual temperature variation as a time machine to understand the effects of long‐term climate change on a poleward range shift

Range shifts due to annual variation in temperature are more tractable than range shifts linked to decadal to century long temperature changes due to climate change, providing natural experiments to determine the mechanisms responsible for driving long‐term distributional shifts. In this study we couple physiologically grounded mechanistic models with biogeographic surveys in 2 years with high levels of annual temperature variation to disentangle the drivers of a historical range shift driven by climate change. The distribution of the barnacle Semibalanus balanoides has shifted 350 km poleward in the past half century along the east coast of the United States. Recruits were present throughout the historical range following the 2015 reproductive season, when temperatures were similar to those in the past century, and absent following the 2016 reproductive season when temperatures were warmer than they have been since 1870, the earliest date for temperature records. Our dispersal dependent mechanistic models of reproductive success were highly accurate and predicted patterns of reproduction success documented in field surveys throughout the historical range in 2015 and 2016. Our mechanistic models of reproductive success not only predicted recruitment dynamics near the range edge but also predicted interior range fragmentation in a number of years between 1870 and 2016. All recruits monitored within the historical range following the 2015 colonization died before 2016 suggesting juvenile survival was likely the primary driver of the historical range retraction. However, if 2016 is indicative of future temperatures mechanisms of range limitation will shift and reproductive failure will lead to further range retraction in the future. Mechanistic models are necessary for accurately predicting the effects of climate change on ranges of species.     

Quantitative losses vs. qualitative stability of ectomycorrhizal community responses to 3 years of experimental summer drought in a beech–spruce forest

Forest ecosystems in central Europe are predicted to face an increasing frequency and severity of summer droughts because of global climate change. European beech and Norway spruce often coexist in these forests with mostly positive effects on their growth. However, their different below‐ground responses to drought may lead to differences in ectomycorrhizal (ECM) fungal community composition and functions which we examined at the individual root and ecosystem levels. We installed retractable roofs over plots in Kranzberg Forest (11°39′42″E, 48°25′12″N; 490 m a.s.l.) to impose repeated summer drought conditions and assigned zones within each plot where trees neighboured the same or different species to study mixed species effects. We found that ECM fungal community composition changed and the numbers of vital mycorrhizae decreased for both tree species over 3 drought years (2014–2016), with the ECM fungal community diversity of beech exhibiting a faster and of spruce a stronger decline. Mixed stands had a positive effect on the ECM fungal community diversity of both tree species after the third drought year. Ectomycorrhizae with long rhizomorphs increased in both species under drought, indicating long‐distance water transport. However, there was a progressive decline in the number of vital fine roots during the experiment, resulting in a strong reduction in enzyme activity per unit volume of soil. Hydrolytic enzyme activities of the surviving ectomycorrhizae were stable or stimulated upon drought, but there was a large decline in ECM fungal species with laccase activity, indicating a decreased potential to exploit nutrients bound to phenolic compounds. Thus, the ectomycorrhizae responded to repeated drought by maintaining or increasing their functionality at the individual root level, but were unable to compensate for quantitative losses at the ecosystem level. These findings demonstrate a strong below‐ground impact of recurrent drought events in forests.