Global variations in ecosystem‐scale isohydricity

Droughts are expected to become more frequent and more intense under climate change. Plant mortality rates and biomass declines in response to drought depend on stomatal and xylem flow regulation. Plants operate on a continuum of xylem and stomatal regulation strategies from very isohydric (strict regulation) to very anisohydric. Coexisting species may display a variety of isohydricity behaviors. As such, it can be difficult to predict how to model the degree of isohydricity at the ecosystem scale by aggregating studies of individual species. This is nonetheless essential for accurate prediction of ecosystem drought resilience. In this study, we define a metric for the degree of isohydricity at the ecosystem scale in analogy with a recent metric introduced at the species level. Using data from the AMSR‐E satellite, this metric is evaluated globally based on diurnal variations in microwave vegetation optical depth (VOD), which is directly related to leaf water potential. Areas with low annual mean radiation are found to be more anisohydric. Except for evergreen broadleaf forests in the tropics, which are very isohydric, and croplands, which are very anisohydric, land cover type is a poor predictor of ecosystem isohydricity, in accordance with previous species‐scale observations. It is therefore also a poor basis for parameterizing water stress response in land‐surface models. For taller ecosystems, canopy height is correlated with higher isohydricity (so that rainforests are mostly isohydric). Highly anisohydric areas show either high or low underlying water use efficiency. In seasonally dry locations, most ecosystems display a more isohydric response (increased stomatal regulation) during the dry season. In several seasonally dry tropical forests, this trend is reversed, as dry‐season leaf‐out appears to coincide with a shift toward more anisohydric strategies. The metric developed in this study allows for detailed investigations of spatial and temporal variations in plant water behavior.     

Conditionally autoregressive models improve occupancy analyses of autocorrelated data: An example with environmental DNA

Site occupancy‐detection models (SODMs) are statistical models widely used for biodiversity surveys where imperfect detection of species occurs. For instance, SODMs are increasingly used to analyse environmental DNA (eDNA) data, taking into account the occurrence of both false‐positive and false‐negative errors. However, species occurrence data are often characterized by spatial and temporal autocorrelation, which might challenge the use of standard SODMs. Here we reviewed the literature of eDNA biodiversity surveys and found that most of studies do not take into account spatial or temporal autocorrelation. We then demonstrated how the analysis of data with spatial or temporal autocorrelation can be improved by using a conditionally autoregressive SODM, and show its application to environmental DNA data. We tested the autoregressive model on both simulated and real data sets, including chronosequences with different degrees of autocorrelation, and a spatial data set on a virtual landscape. Analyses of simulated data showed that autoregressive SODMs perform better than traditional SODMs in the estimation of key parameters such as true‐/false‐positive rates and show a better discrimination capacity (e.g., higher true skill statistics). The usefulness of autoregressive SODMs was particularly high in data sets with strong autocorrelation. When applied to real eDNA data sets (eDNA from lake sediment cores and freshwater), autoregressive SODM provided more precise estimation of true‐/false‐positive rates, resulting in more reasonable inference of occupancy states. Our results suggest that analyses of occurrence data, such as many applications of eDNA, can be largely improved by applying conditionally autoregressive specifications to SODMs.     

Short‐term effects of a severe drought on avian diversity and abundance in a Pampas Agroecosystem

Agroecosystems are naturally variable at different scales showing strong environmental variations through time and in space. Therefore, temporal dynamics should be taken into account to understand the species‐habitat relationship and provide information for biodiversity management. Droughts are climatic events that introduce variations in environmental conditions by reducing food resources and are increasing in severity and frequency due to global climate change. In 2008, a severe drought occurred in the argentine Pampas, which allowed us to test its short‐term effect on avian diversity patterns according to different land uses. Also, we could test how variations in net primary productivity affected bird populations and explore the usefulness of this ecological process as an indicator of ecosystem service supply applicable to different ecological contexts. We surveyed bird diversity in two consecutive years (2007–2008) in the Rolling Pampas of Argentina, and we assessed environmental attributes at two spatial scales. We explored the relationship between bird diversity and an ecosystem services provision index (ESPI) that uses a proxy of net primary productivity and its intra‐annual variation to test its adequacy in the changing conditions, we conducted our surveys. Results showed that drought affected negatively both species richness and abundance as a consequence of net primary productivity reductions. There was not a clear association of diversity changes with land use, and it is probable that the effect of drought is a complex combination of productivity, land use and spatial scale. ESPI proved robust in front of the environmental changes, and its predictive capacity was better at larger scales. These results are promising for the assessment of ecosystem services provision in a context of global climate changes. Abstract in Spanish is available with online material.     

Environmental heterogeneity and spatiotemporal variability in plant defense traits

At the scale of the local plant community, we know very little about how spatial and temporal environmental heterogeneity affects the diversity in types and levels of plant defenses. If environmental heterogeneity is an important mechanism influencing plant defense traits, then defense expression should co‐vary spatially with environmental conditions and change as succession progresses. In this study, we examined how spatial heterogeneity and succession influence putative resistance and tolerance trait levels in late goldenrod Solidago altissima. We quantified the spatial distributions of herbivore damage and traits associated with resistance (leaf toughness, phenolics), tolerance (specific leaf area, relative growth rate, leaf addition rate and leaf senescence rate), and fitness (height, diameter, inflorescence biomass) of goldenrods within replicate early‐ and late‐successional fields. Also, we characterized the local neighborhood (stem density, canopy cover, ground vegetative cover) and edaphic conditions (soil moisture, pH, N) surrounding each target ramet, and determined relationships between these environmental variables and goldenrod trait levels. The distribution of traits within fields was strongly non‐random, and defense‐trait levels were more strongly spatially structured (i.e. autocorrelated) in late‐ than in early‐successional fields. Also, defense traits were most strongly correlated with aspects of the local plant neighborhood, and these relationships differed in important ways between successional stages. In late‐successional fields, tolerance trait specific leaf area was positively correlated with canopy cover and negatively correlated with stem density. In early‐successional fields, the relationship between ground vegetative cover and resistance (i.e. 1 – damage) was significantly stronger than in late‐successional fields. A novel insight from this study is the possibility that changes in the biotic environment during succession may shift the expression of defense from a resistance to a tolerance strategy in our system. This study highlights the context dependence of plant defense trait levels, which may promote their spatial and temporal variability in heterogeneous landscapes.     

中国遥感干旱指数时空特征及其对气候和地表覆盖变化的响应

干旱严重影响植被生长,威胁粮食安全,基于遥感计算的植被状态指数（Vegetation Condition Index,VCI）、温度状态指数（Temperature Condition Index,TCI）和植被健康指数（Vegetation Health Index,VHI）是常用的干旱指数,被广泛应用于干旱监测。为了探究近年来我国干旱特征及其对气候和地表覆盖变化的响应,分析了2003-2016年期间VCI、TCI和VHI的时空变化特征;采用最小二乘（OLS）和偏相关分析方法分析了这些指数对气候和地表覆盖变化的响应。基于上述干旱指数计算的干旱频率表明,中温带中部和南温带等地区干旱发生频率高,干旱指数变化趋势表明在2003-2016年期间中国大部分地区干旱缓解,但在中温带、南温带和高原气候区等局部地区干旱加剧;总体而言,干旱指数随着年平均温度的上升和年降水量的降低而减小,VHI与温度和降水量的相关性在不同气候区的一致性优于VCI和TCI;裸土的减少和植被的增加导致干旱指数增大,树木转变为低矮植被干旱指数降低。     

Availability of food resources and habitat structure shape the individual‐resource network of a Neotropical marsupial

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Range expansion through fragmented landscapes under a variable climate

Ecological responses to climate change may depend on complex patterns of variability in weather and local microclimate that overlay global increases in mean temperature. Here, we show that high‐resolution temporal and spatial variability in temperature drives the dynamics of range expansion for an exemplar species, the butterfly Hesperia comma. Using fine‐resolution (5 m) models of vegetation surface microclimate, we estimate the thermal suitability of 906 habitat patches at the species' range margin for 27 years. Population and metapopulation models that incorporate this dynamic microclimate surface improve predictions of observed annual changes to population density and patch occupancy dynamics during the species' range expansion from 1982 to 2009. Our findings reveal how fine‐scale, short‐term environmental variability drives rates and patterns of range expansion through spatially localised, intermittent episodes of expansion and contraction. Incorporating dynamic microclimates can thus improve models of species range shifts at spatial and temporal scales relevant to conservation interventions.     

Scales of spatiotemporal variability in macroinvertebrate abundance and diversity in monsoonal streams: detecting environmental change

1. We quantified spatial and temporal variability in benthic macroinvertebrate species richness, diversity and abundance in six unpolluted streams in monsoonal Hong Kong at different scales using a nested sampling design. The spatial scales were regions, stream sites and stream sections within sites; temporal scales were years (1997–99), seasons (dry versus wet seasons) and days within seasons. 2. Spatiotemporal variability in total abundance and species richness was greater during the wet season, especially at small scales, and tended to obscure site‐ and region‐scale differences, which were more conspicuous during the dry season. Total abundance and richness were greater in the dry season, reflecting the effects of spate‐induced disturbance during the wet season. Species diversity showed little variation at the seasonal scale, but variability at the site scale was apparent during both seasons. 3. Despite marked variations in monsoonal rainfall, inter‐year differences in macroinvertebrate richness and abundance at the site scale during the wet season were minor. Inter‐year differences were only evident during the dry season when streams were at base flow and biotic interactions may structure assemblages. 4. Small‐scale patchiness within riffles was the dominant spatial scale of variation in macroinvertebrate richness, total abundance and densities of common species, although site or region was important for some species. The proportion of total variance contributed by small‐scale spatial variability increased during the dry season, whereas temporal variability associated with days was greater during the wet season. 5. The observed patterns of spatiotemporal variation have implications for detection of environmental change or biomonitoring using macroinvertebrate indicators in streams in monsoonal regions. Sampling should be confined to the dry season or, in cases where more resources are available, make use of data from both dry and wet seasons. Sampling in more than one dry season is required to avoid the potentially confounding effects of inter‐year variation, although variability at that scale was relatively small.     

Water as a resource,stress and disturbance shaping tundra vegetation

Water is crucial for plant productivity and survival as a fundamental resource, but water conditions can also cause physiological stress and mechanical disturbance to vegetation. However, these different influences of water on vegetation patterns have not been evaluated simultaneously. Here, we demonstrate the importance of three water aspects (spatial and temporal variation of soil moisture and fluvial disturbance) for three ecologically and evolutionary distinct taxonomical groups (vascular plants, mosses and lichens) in Fennoscandian mountain tundra. Fine‐scale plant occurrence data for 271 species were collected from 378 × 1 m² plots sampled over broad environmental gradients (water, temperature, radiation, soil pH, cryogenic processes and the dominant allelopathic plant species). While controlling all other key environmental variables, water in its different aspects proved to be a crucial environmental driver, acting on individual species and on community characteristics. The inclusion of the water variables significantly improved our models. In this high‐latitude system, the importance of spatial variability of water exceeds the importance of temperature for the fine‐scale distribution of species from the three taxonomical groups. We found differing responses to the three water variables between and within the taxonomical groups. Water as a resource was the most important water‐related variable in species distribution models across all taxonomical groups. Both water resource and disturbance were strongly related to vascular plant species richness, whereas for moss species richness, water resources had the highest influence. For lichen species richness, water disturbance was the most influential water‐related variable. These findings demonstrate that water variables are not only independent properties of tundra hydrology, but also that water is truly a multifaceted driver of vegetation patterns at high‐latitudes.     

Demographic response of plant populations to habitat fragmentation and temporal environmental variability

Many plant species currently exist in fragmented populations of different sizes, while they also experience unpredictable climatic fluctuation over time. However, we still understand little about how plant demography responds to such spatial and temporal environmental variability. We studied population dynamics of an understory perennial herb Trillium camschatcense in the Tokachi plain of Hokkaido, Japan, where a significant effect of forest fragmentation on seedling recruitment was previously reported. Four populations across a range of fragment sizes were studied for 6 years, and the data were analyzed using matrix population models. Per capita fecundity (the number of recruits per plant) varied greatly among populations, but the variation in population growth rates (λ) was mainly driven by the variation in stasis and growth rates, suggesting that the general trend of reduced fecundity in fragmented populations may not be readily translated into subsequent dynamics. Temporal variation in λ among years was more than 2 times larger than spatial variation among populations, and this result was likely attributable to the contrasting response of correlation structures among demographic rates. The among-population variation in λ was dampened by negative covariation between matrix elements possibly due to density-dependent regulation as well as an inherent constraint that some elements are not independent, whereas positive covariation between matrix elements resulted in large temporal variation in λ. Our results show that population dynamics responded differently to habitat fragmentation and temporal variability of the environment, emphasizing the need to discriminate these spatial and temporal variations in demographic models. Although no populations were projected to be declining in stochastic simulations, correlation between current habitat size and plant density implies historical λ is positively related to habitat size.     

Environmental drivers interactively affect individual tree growth across temperate European forests

Forecasting the growth of tree species to future environmental changes requires a better understanding of its determinants. Tree growth is known to respond to global‐change drivers such as climate change or atmospheric deposition, as well as to local land‐use drivers such as forest management. Yet, large geographical scale studies examining interactive growth responses to multiple global‐change drivers are relatively scarce and rarely consider management effects. Here, we assessed the interactive effects of three global‐change drivers (temperature, precipitation and nitrogen deposition) on individual tree growth of three study species (Quercus robur/petraea, Fagus sylvatica and Fraxinus excelsior). We sampled trees along spatial environmental gradients across Europe and accounted for the effects of management for Quercus. We collected increment cores from 267 trees distributed over 151 plots in 19 forest regions and characterized their neighbouring environment to take into account potentially confounding factors such as tree size, competition, soil conditions and elevation. We demonstrate that growth responds interactively to global‐change drivers, with species‐specific sensitivities to the combined factors. Simultaneously high levels of precipitation and deposition benefited Fraxinus, but negatively affected Quercus’ growth, highlighting species‐specific interactive tree growth responses to combined drivers. For Fagus, a stronger growth response to higher temperatures was found when precipitation was also higher, illustrating the potential negative effects of drought stress under warming for this species. Furthermore, we show that past forest management can modulate the effects of changing temperatures on Quercus’ growth; individuals in plots with a coppicing history showed stronger growth responses to higher temperatures. Overall, our findings highlight how tree growth can be interactively determined by global‐change drivers, and how these growth responses might be modulated by past forest management. By showing future growth changes for scenarios of environmental change, we stress the importance of considering multiple drivers, including past management and their interactions, when predicting tree growth.     

When can refuges mediate the genetic effects of fire regimes? A simulation study of the effects of topography and weather on neutral and adaptive genetic diversity in fire‐prone landscapes

Understanding how landscape heterogeneity mediates the effects of fire on biodiversity is increasingly important under global changes in fire regimes. We used a simulation experiment to investigate how fire regimes interact with topography and weather to shape neutral and selection‐driven genetic diversity under alternative dispersal scenarios, and to explore the conditions under which microrefuges can maintain genetic diversity of populations exposed to recurrent fire. Spatial heterogeneity in simulated fire frequency occurred in topographically complex landscapes, with fire refuges and fire‐prone “hotspots” apparent. Interannual weather variability reduced the effect of topography on fire patterns, with refuges less apparent under high weather variability. Neutral genetic diversity was correlated with long‐term fire frequency under spatially heterogeneous fire regimes, being higher in fire refuges than fire‐prone areas, except under high dispersal or low fire severity (low mortality). This generated different spatial genetic structures in fire‐prone and fire‐refuge components of the landscape, despite similar dispersal. In contrast, genetic diversity was only associated with time since the most recent fire in flat landscapes without predictable refuges and hotspots. Genetic effects of selection driven by fire‐related conditions depended on selection pressure, migration distance and spatial heterogeneity in fire regimes. Allele frequencies at a locus conferring higher fitness under successional environmental conditions followed a pattern of “temporal adaptation” to contemporary conditions under strong selection pressure and high migration. However, selected allele frequencies were correlated with spatial variation in long‐term mean fire frequency (relating to environmental predictability) under weak dispersal, low selection pressure and strong spatial heterogeneity in fire regimes.     

Disentangling environmental and spatial processes of community assembly in tropical forests from local to regional scales

Understanding patterns and mechanisms of variation in the compositional structure of communities across spatial scales is one of the fundamental challenges in ecology and biogeography. In this study, we evaluated the effects of spatial extent (i.e. size of study region) on: 1) whether community composition can be better explained by environmental (i.e. niche‐based) or spatial (e.g. dispersal‐based) processes ; and 2) how climate and soils contribute to the influence of environment on plant community composition. We surveyed community composition across a network of 398 forest plots spanning a ～4000 m elevational gradient in the Madidi region in northwestern Bolivia. Using redundancy analyses and hierarchical variation partitioning, we disentangled the effects of environmental and spatial predictors on species composition, further decomposing the environmental effect between its climatic and soil components. We repeated analyses for 200 sub‐regions ranging in spatial extent from ～250 to ～17 500 km². Our analyses show a high degree of idiosyncrasy in results that come from different sub‐regions. Despite this variability, we were able to identify various important patterns in the structure of tropical plant communities in our study system. First, even though sub‐regions varied in size by nearly two orders of magnitude, the total amount of explained variation in community composition was scale independent; at all spatial scales, environment and space accounted for about 25% of the differences in community composition among plots. Second, the measured environmental effect was higher than the spatial effect on average and in the vast majority of sub‐regions. This was true regardless of the spatial extent of analysis. Finally, we found that both climatic and soil variables accounted for significant fractions of variation, but climate was always more important than soils.     

Beyond soil water potential: An expanded view on isohydricity including land–atmosphere interactions and phenology

Over the past decade, the concept of isohydry or anisohydry, which describes the link between soil water potential (Ψ_S), leaf water potential (Ψ_L), and stomatal conductance (g_s), has soared in popularity. However, its utility has recently been questioned, and a surprising lack of coordination between the dynamics of Ψ_L and g_s across biomes has been reported. Here, we offer a more expanded view of the isohydricity concept that considers effects of vapour pressure deficit (VPD) and leaf area index (A_L) on the apparent sensitivities of Ψ_L and g_s to drought. After validating the model with tree‐ and ecosystem‐scale data, we find that within a site, isohydricity is a strong predictor of limitations to stomatal function, though variation in VPD and leaf area, among other factors, can challenge its diagnosis. Across sites, the theory predicts that the degree of isohydricity is a good predictor of the sensitivity of g_s to declining soil water in the absence of confounding effects from other drivers. However, if VPD effects are significant, they alone are sufficient to decouple the dynamics of Ψ_L and g_s entirely. We conclude with a set of practical recommendations for future applications of the isohydricity framework within and across sites.     

Interspecific competition alters leaf stoichiometry in 20 grassland species

The extensive use of traits in ecological studies over the last few decades to predict community functions has revealed that plant traits are plastic and respond to various environmental factors. These plant traits are assumed to predict how plants compete and capture resources. Variation in stoichiometric ratios both within and across species reflects resource capture dynamics under competition. However, the impact of local plant diversity on species‐specific stoichiometry remains poorly studied. Here, we analyze how spatial and temporal diversity in resource‐acquisition traits affects leaf elemental stoichiometry of plants (i.e. the result of resource capture) and how flexible this stoichiometry is depending on the functional composition of the surrounding community. Therefore, we assessed inter‐ and intraspecific variations of leaf carbon (C), nitrogen (N), and phosphorus (P) (and their ratios) of 20 grassland species in a large trait‐based plant diversity experiment located in Jena (Germany) by measuring leaf elemental concentrations at the species‐level along a gradient in plant trait dissimilarity. Our results show that plants showed large intra‐ and interspecific variation in leaf stoichiometry, which was only partly explained by the functional group identity (grass or herb) of the species. Elemental concentrations (N, P, but not C) decreased with plant species richness, and species tended to become more deviant from their monoculture stoichiometry with increasing trait dissimilarity in the community. These responses differed among species, some consistently increased or decreased in P and N concentrations; for other species, the negative or positive change in P and N concentrations increased with increasing trait difference between the target species and the remaining community. The strength of this relationship was significantly associated to the relative position of the species along trait gradients related to resource acquisition. Trait‐difference and trait‐diversity thus were important predictors of how species’ resource capture changed in competitive neighbourhoods.     

Lifelong foraging and individual specialisation are influenced by temporal changes of resource availability

Resource availability largely determines the distribution and behaviour of organisms. In plant–pollinator communities, availability of floral resources may change so rapidly that pollinator individuals can benefit from switching between multiple resources, i.e. different flowering plant species. Insect pollinator individuals of a given generation often occur in different time windows during the reproductive season. This temporal variation in individual occurrences, together with the rapidly changing resource availability, may lead individuals of the same population to encounter and use different resources, resulting in an apparent individual specialisation. We hypothesized, that 1) individual pollinators change their resource use (flower visitation) during their lifetime according to the changing availability of floral resources, and that 2) temporal variation in individual occurrences of pollinators and in resource availability will partly explain individual specialisation. To test these hypotheses, we observed flower visitations of individually marked clouded Apollo butterflies Parnassius mnemosyne during one reproductive season. We found temporal changes in lifetime individual resource use that followed the changes in resource availability, indicating that butterflies can adjust foraging to varying resource availability. Individuals differed considerably in their resource use. This variation was partly explained by temporal variation in both floral resource availability and temporal occurrence of individual butterflies. We suggest the butterfly as a sequential specialist, i.e. short‐term specialist and long‐term generalist. This foraging plasticity can be essential for short‐living insect pollinators in rapidly changing environments. Although flowering dynamics do not fully explain the variability in foraging, our results highlight the importance of temporal dimension in resource use studies. Ultimately, the relative pace of environmental change compared to individual lifespan may be a key factor in resource use plasticity.     

Characterizing habitat suitability for a central‐place forager in a dynamic marine environment

Characterizing habitat suitability for a marine predator requires an understanding of the environmental heterogeneity and variability over the range in which a population moves during a particular life cycle. Female California sea lions (Zalophus californianus) are central‐place foragers and are particularly constrained while provisioning their young. During this time, habitat selection is a function of prey availability and proximity to the rookery, which has important implications for reproductive and population success. We explore how lactating females may select habitat and respond to environmental variability over broad spatial and temporal scales within the California Current System. We combine near‐real‐time remotely sensed satellite oceanography, animal tracking data (n = 72) from November to February over multiple years (2003–2009) and Generalized Additive Mixed Models (GAMMs) to determine the probability of sea lion occurrence based on environmental covariates. Results indicate that sea lion presence is associated with cool ( <14°C ), productive waters, shallow depths, increased eddy activity, and positive sea‐level anomalies. Predictive habitat maps generated from these biophysical associations suggest winter foraging areas are spatially consistent in the nearshore and offshore environments, except during the 2004–2005 winter, which coincided with an El Niño event. Here, we show how a species distribution model can provide broadscale information on the distribution of female California sea lions during an important life history stage and its implications for population dynamics and spatial management.     

Management defines species turnover of bees and flowering plants in vineyards

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Predator Assemblage Structure and Temporal Variability of Species Richness and Abundance in Forests of High Tree Diversity

Predators significantly affect ecosystem functions, but our understanding of to what extent findings can be transferred from experiments and low‐diversity systems to highly diverse, natural ecosystems is limited. With a particular threat of biodiversity loss at higher trophic levels, however, knowledge of spatial and temporal patterns in predator assemblages and their interrelations with lower trophic levels is essential for assessing effects of trophic interactions and advancing biodiversity conservation in these ecosystems. We analyzed spatial and temporal variability of spider assemblages in tree species‐rich subtropical forests in China, across 27 study plots varying in woody plant diversity and stand age. Despite effects of woody plant richness on spider assemblage structure, neither habitat specificity nor temporal variability of spider richness and abundance were influenced. Rather, variability increased with forest age, probably related to successional changes in spider assemblages. Our results indicate that woody plant richness and theory predicting increasing predator diversity with increasing plant diversity do not necessarily play a major role for spatial and temporal dynamics of predator assemblages in such plant species‐rich forests. Diversity effects on biotic or abiotic habitat conditions might be less pronounced across our gradient from medium to high plant diversity than in previously studied less diverse systems, and bottom‐up effects might level out at high plant diversity. Instead, our study highlights the importance of overall (diversity‐independent) environmental heterogeneity in shaping spider assemblages and, as indicated by a high species turnover between plots, as a crucial factor for biodiversity conservation at a regional scale in these subtropical forests.