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Aim: Isotrema is a monophyletic subgenus of Aristolochia with species in East Asia and North/Central America. Earlier studies, based on limited sampling, suggested that the Asian and American species do not form two reciprocal sister clades. We reconstructed phylogenetic relationships within Isotrema to infer the biogeographical events that have shaped its present-day distribution. Location: Eastern Asia, North America, Mexico, Central America. Methods: We performed parsimony and Bayesian phylogenetic analyses of 54 accessions using three chloroplast regions. The temporal origins were traced with relaxed phylogenetics and penalized likelihood using fossil calibrations; these methods were combined with ancestral area reconstructions in a comparative approach using statistical dispersal-vicariance analysis (S-DIVA) and dispersal-extinction-cladogenesis (DEC) analyses. Results: The ancestors of the herbaceous eastern North American species and the woody species probably diverged during the Oligocene. The woody species form a Neotropical and a North American/Eastern Asian clade. Diversification in the Neotropical and North American/Eastern Asian clades occurred mostly in the Miocene. In the latter clade, two further intercontinental splits are reconstructed: between the exclusively western North American Aristolochia californica and most of the Asian taxa, and between A. manshuriensis and A. tomentosa. Main conclusions : The present distribution of Isotrema developed via a number of dispersal, vicariance and extinction events. The disjunct distributions observed may be the result mainly of non-synchronous events (e.g. a decrease of mean annual temperature in the Oligocene and the development of unfavourable conditions across the Bering land bridge) that were responsible for the fragmentation of the mesophytic forests. Later diversifications may be correlated with events such as the orogeny of the Western Cordillera and Appalachian Mountains, the development of extensive grasslands in North America, and the mainland extension of Central America southwards to western and central Panama

1. The subdiscipline of ‘community phylogenetics’ is rapidly growing and influencing thinking regarding community assembly. In particular, phylogenetic dispersion of co-occurring species within a community is commonly used as a proxy to identify which community assembly processes may have structured a particular community: phylogenetic clustering as a proxy for abiotic assembly, that is habitat filtering, and phylogenetic overdispersion as a proxy for biotic assembly, notably competition.2. We challenge this approach by highlighting (typically) implicit assumptions that are, in reality, only weakly supported, including (i) phylogenetic dispersion reflects trait dispersion; (ii) a given ecological function can be performed only by a single trait state or combination of trait states; (iii)trait similarity causes enhanced competition; (iv) competition causes species exclusion; (v) communities are at equilibrium with processes of assembly having been completed; (vi) assembly through habitat filtering decreases in importance if assembly through competition increases, such that the relative balance of the two can be thus quantified by a single parameter; and (vii) observed phylogenetic dispersion is driven predominantly by local and present-day processes.3. Moreover, technical sophistication of the phylogenetic-patterns-as-proxy approach trades off against sophistication in alternative, potentially more pertinent approaches to directly observe or manipulate assembly processes.4. Despite concerns about using phylogenetic dispersion as a proxy for community assembly processes, we suggest there are underappreciated benefits of quantifying the phylogenetic structure of communities, including (i) understanding how coexistence leads to the macroevolutionary diversification of habitat lineage-pools (i.e. phylogenetic-patterns-as-result approach); and (ii) understanding the macroevolutionary contingency of habitat lineage-pools and how it affects present-day species coexistence in local communities (i.e. phylogeneticpatterns- as-cause approach).5. We conclude that phylogenetic patterns may be little useful as proxy of community assembly. However, such patterns can prove useful to identify and test novel hypotheses on (i) how local coexistence may control macroevolution of the habitat lineage-pool, for example through competition among close relatives triggering displacement and diversification of characters, and (ii) how macroevolution within the habitat lineage-pool may control local coexistence of related species, for example through origin of close relatives that can potentially enter in competition.

Collinearity refers to the non independence of predictor variables, usually in a regression-type analysis. It is a common feature of any descriptive ecological data set and can be a problem for parameter estimation because it inflates the variance of regression parameters and hence potentially leads to the wrong identification of relevant predictors in a statistical model. Collinearity is a severe problem when a model is trained on data from one region or time, and predicted to another with a different or unknown structure of collinearity. To demonstrate the reach of the problem of collinearity in ecology, we show how relationships among predictors differ between biomes, change over spatial scales and through time. Across disciplines, different approaches to addressing collinearity problems have been developed, ranging from clustering of predictors, threshold-based pre-selection, through latent variable methods, to shrinkage and regularisation. Using simulated data with five predictor-response relationships of increasing complexity and eight levels of collinearity we compared ways to address collinearity with standard multiple regression and machine-learning approaches. We assessed the performance of each approach by testing its impact on prediction to new data. In the extreme, we tested whether the methods were able to identify the true underlying relationship in a training dataset with strong collinearity by evaluating its performance on a test dataset without any collinearity. We found that methods specifically designed for collinearity, such as latent variable methods and tree based models, did not outperform the traditional GLM and threshold-based pre-selection. Our results highlight the value of GLM in combination with penalised methods (particularly ridge) and threshold-based pre-selection when omitted variables are considered in the final interpretation. However, all approaches tested yielded degraded predictions under change in collinearity structure and the folk lore'-thresholds of correlation coefficients between predictor variables of |r| >0.7 was an appropriate indicator for when collinearity begins to severely distort model estimation and subsequent prediction. The use of ecological understanding of the system in pre-analysis variable selection and the choice of the least sensitive statistical approaches reduce the problems of collinearity, but cannot ultimately solve them.

Mixing of complementary tree species may increase stand productivity, mitigate the effects of drought and other risks, and pave the way to forest production systems which may be more resource-use efficient and stable in the face of climate change. However, systematic empirical studies on mixing effects are still missing for many commercially important and widespread species combinations. Here we studied the growth of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) in mixed versus pure stands on 32 triplets located along a productivity gradient through Europe, reaching from Sweden to Bulgaria and from Spain to the Ukraine. Stand inventory and taking increment cores on the mainly 60-80 year-old trees and 0.02-1.55 ha sized, fully stocked plots provided insight how species mixing modifies the structure, dynamics and productivity compared with neighbouring pure stands. In mixture standing volume (+12 %), stand density (+20 %), basal area growth (+12 %), and stand volume growth (+8 %) were higher than the weighted mean of the neighbouring pure stands. Scots pine and European beech contributed rather equally to the overyielding and overdensity. In mixed stands mean diameter (+20 %) and height (+6 %) of Scots pine was ahead, while both diameter and height growth of European beech were behind (-8 %). The overyielding and overdensity were independent of the site index, the stand growth and yield, and climatic variables despite the wide variation in precipitation (520-1175 mm year(-1)), mean annual temperature (6-10.5 A degrees C), and the drought index by de Martonne (28-61 mm A degrees C-1) on the sites. Therefore, this species combination is potentially useful for increasing productivity across a wide range of site and climatic conditions. Given the significant overyielding of stand basal area growth but the absence of any relationship with site index and climatic variables, we hypothesize that the overyielding and overdensity results from several different types of interactions (light-, water-, and nutrient-related) that are all important in different circumstances. We discuss the relevance of the results for ecological theory and for the ongoing silvicultural transition from pure to mixed stands and their adaptation to climate change.

In this review, the unique features and facts of long-term experiments are presented. Long-term experimental plots provide information of forest stand dynamics which cannot be derived from forest inventories or small temporary plots. Most comprise unthinned plots which represent the site specific maximum stand density as an unambiguous reference. By measuring the remaining as well as the removed stand, the survey of long-term experiments provides the total production at a given site, which is most relevant for examining the relationship between site conditions and stand productivity on the one hand and between stand density and productivity on the other. Thus, long-term experiments can reveal the site-specific effect of thinning and species mixing on stand structure, production and carbon sequestration. If they cover an entire rotation or even the previous and following generation on a given site, they reveal a species’ long-term behaviour and any growth trends caused by environmental changes. Second, we exploit the unique data of European long-term experiments, some of which have been surveyed since 1848. We show the long-term effect of different density regimes on stand dynamics and an essential trade-off between total stand volume production and mean tree size. Long-term experiments reveal that tree species mixing can significantly increase stand density and productivity compared with monospecific stands. Thanks to surveys spanning decades or even a century, we can show the changing long-term-performance of different provenances and acceleration of stand production caused by environmental change, as well as better understand the growth dynamics of natural forests. Without long-term experiments forest science and practice would be not in a position to obtain such findings which are of the utmost relevance for science and practice. Third, we draw conclusions and show perspectives regarding the maintenance and further development of long-term experiments. It would require another 150 years to build up a comparable wealth of scientific information, practical knowledge, and teaching and training model examples. Although tempting, long-term experiments should not be sacrificed for cost-cutting measures. Given the global environmental change and the resulting challenges for sustainable management, the network of long-term experiments should rather be extended regarding experimental factors, recorded variables and inter- and transdisciplinary use for science and practice.

The essential mineral nutrient potassium (K(+)) is the most important inorganic cation for plants and is recognized as a limiting factor for crop yield and quality. Nonetheless, it is only partially understood how K(+) contributes to plant productivity. K(+) is used as a major active solute to maintain turgor and to drive irreversible and reversible changes in cell volume. K(+) also plays an important role in numerous metabolic processes, for example, by serving as an essential cofactor of enzymes. Here, we provide evidence for an additional, previously unrecognized role of K(+) in plant growth. By combining diverse experimental approaches with computational cell simulation, we show that K(+) circulating in the phloem serves as a decentralized energy storage that can be used to overcome local energy limitations. Posttranslational modification of the phloem-expressed Arabidopsis K(+) channel AKT2 taps this \"potassium battery,\" which then efficiently assists the plasma membrane H(+)-ATPase in energizing the transmembrane phloem (re)loading processes.

Background Loss of biodiversity and increased nutrient inputs are two of the most crucial anthropogenic factors driving ecosystem change. Although both received considerable attention in previous studies, information on their interactive effects on ecosystem functioning is scarce. In particular, little is known on how soil biota and their functions are affected by combined changes in plant diversity and fertilization. Methodology/Principal Findings We investigated the effects of plant diversity, functional community composition, and fertilization on the biomass and respiration of soil microbial communities in a long-term biodiversity experiment in semi-natural grassland (Jena Experiment). Plant species richness enhanced microbial basal respiration and microbial biomass, but did not significantly affect microbial specific respiration. In contrast, the presence of legumes and fertilization significantly decreased microbial specific respiration, without altering microbial biomass. The effect of legumes was superimposed by fertilization as indicated by a significant interaction between the presence of legumes and fertilization. Further, changes in microbial stoichiometry (C-to-N ratio) and specific respiration suggest the presence of legumes to reduce N limitation of soil microorganisms and to modify microbial C use efficiency. Conclusions/Significance Our study highlights the role of plant species and functional group diversity as well as interactions between plant community composition and fertilizer application for soil microbial functions. Our results suggest soil microbial stoichiometry to be a powerful indicator of microbial functioning under N limited conditions. Although our results support the notion that plant diversity and fertilizer application independently affect microbial functioning, legume effects on microbial N limitation were superimposed by fertilization, indicating significant interactions between the functional composition of plant communities and nutrient inputs for soil processes.

The spatiotemporal dynamics of, and interspecific differences in, the acquisition of litter-derived nitrogen (N) by natural assemblages of ectomycorrhizal root tips are poorly understood. Small cylindrical mesh bags containing 15N-labelled beech (Fagus sylvatica) leaf litter that permit hyphal but not root ingrowth were inserted vertically into the top soil layer of an old-growth beech forest. The lateral transfer of 15N into the circumjacent soil, roots, microbes and ectomycorrhizas was measured during an 18-month exposure period. Ectomycorrhial fungi (EMF) showed large interspecific variation in the temporal pattern and extent of 15N accumulation. Initially, when N was mainly available from the leachate, microbes were more efficient at N immobilization than the majority of EMF, but distinct fungal species also showed significant 15N accumulation. During later phases, the enrichment of 15N in Tomentella badia was higher than in microbes and other EMF species. Roots and soil accumulated 15N with a large delay compared with microbes and EMF. Because approximately half of the studied fungal species had direct access to N from leaf litter and the remainder to N from leached compounds, we suggest that EMF diversity facilitates the N utilization of the host by capturing N originating from early-released solutes and late degradation products from a recalcitrant source.

Observations of vegetation-atmosphere exchange of carbon dioxide (CO2) by the eddy covariance (EC) technique are limited by difficult conditions such as nighttime and heterogeneous terrain. Thus, advective flux components are included into the net ecosystem exchange (NEE) budget. However, advection measurements are experimentally challenging and do not always help to solve the night flux problem of the EC technique. This study investigates alternative methods for the observation of horizontal advection, in particular horizontal concentration gradients, as well as different approaches to coordinate rotation and vertical advection. Continuous high-frequency measurements of the horizontal CO2 concentration field are employed and compared to the often used discontinuous sequential sampling. Significant differences were found in the case of 30-min mean concentration values between the conventional discontinuous sampling approach and the complete observation of the time series by continuous sampling. Estimates of vertical advection rely on accurate estimates of vertical wind velocity (). Therefore, different approaches to the planar fit coordinate rotation have been investigated. Sector-wise rotation was able to eliminate directional dependencies of mean . Furthermore, the effect of the data set length used for rotation (window length) was investigated and was found to have significant impact on estimates of vertical advection, with larger window lengths yielding about 50% larger vertical advection. A sequential planar fit with controlled window length is proposed to give reproducible results. The different approaches to the measurement and calculation of horizontal and vertical advection presented are applied to data obtained during the exchange processes in mountainous region experiment at the FLUXNET site Waldstein-Weidenbrunnen (DE-Bay). Estimates of NEE including advection are compared to NEE from turbulent and storage flux alone without advection. NEE including vertical advection with sector-wise planar fit rotation and controlled window length and including horizontal advection from continuous gradient measurements, which were comprehensively bias corrected by a new approach, did compare well with the expected night flux error, with meteorological drivers of the fluxes and with soil chamber measurements. Unrealistically large and noisy values of horizontal advection from the conventional discontinuous sampling approach, which lead to unrealistic values of NEE, could be eliminated by the alternative approaches presented. We therefore suggest the further testing of those approaches at other sites in order to improve the accuracy of advection measurements and, subsequently, estimates of NEE.

The speed and dynamics of range expansions shape species distributions and community composition. Despite the critical impact of population growth rates for range expansion, they are neglected in existing empirical studies, which focus on the investigation of selected life-history traits. Here, we present an approach based on non-invasive genetic capture–mark–recapture data for the estimation of adult survival, fecundity and juvenile survival, which determine population growth. We demonstrate the reliability of our method with simulated data, and use it to investigate life-history changes associated with range expansion in 35 colonies of the bat species Rhinolophus hipposideros. Comparing the demographic parameters inferred for 19 of those colonies which belong to an expanding population with those inferred for the remaining 16 colonies from a non-expanding population reveals that range expansion is associated with higher net reproduction. Juvenile survival was the main driver of the observed reproduction increase in this long-lived bat species with low per capita annual reproductive output. The higher average growth rate in the expanding population was not associated with a trade-off between increased reproduction and survival, suggesting that the observed increase in reproduction stems from a higher resource acquisition in the expanding population. Environmental conditions in the novel habitat hence seem to have an important influence on range expansion dynamics, and warrant further investigation for the management of range expansion in both native and invasive species.