Long-term shift towards shady and nutrient-rich habitats in Central European temperate forests.

Biodiversity world-wide has been under increasing anthropogenic pressure in the past century. The long-term response of biotic communities has been tackled primarily by focusing on species richness, community composition and functionality. Equally important are shifts between entire communities and habitat types, which remain an unexplored level of biodiversity change. We have resurveyed > 2000 vegetation plots in temperate forests in central Europe to capture changes over an average of five decades. The plots were assigned to eight broad forest habitat types using an algorithmic classification system. We analysed transitions between the habitat types and interpreted the trend in terms of changes in environmental conditions. We identified a directional shift along the combined gradients of canopy openness and soil nutrients. Nutrient-poor open-canopy forest habitats have declined strongly in favour of fertile closed-canopy habitats. However, the shift was not uniform across the whole gradients. We conclude that the shifts in habitat types represent a century-long successional trend with significant consequences for forest biodiversity. Open forest habitats should be urgently targeted for plant diversity restoration through the implementation of active management. The approach presented here can be applied to other habitat types and at different spatio-temporal scales.

Projected climate and canopy change lead to thermophilization and homogenization of forest floor vegetation in a hotspot of plant species richness.

Mountain forests are plant diversity hotspots, but changing climate and increasing forest disturbances will likely lead to far-reaching plant community change. Projecting future change, however, is challenging for forest understory plants, which respond to forest structure and composition as well as climate. Here, we jointly assessed the effects of both climate and forest change, including wind and bark beetle disturbances, using the process-based simulation model iLand in a protected landscape in the northern Alps (Berchtesgaden National Park, Germany), asking: (1) How do understory plant communities respond to 21st-century change in a topographically complex mountain landscape, representing a hotspot of plant species richness? (2) How important are climatic changes (i.e., direct climate effects) versus forest structure and composition changes (i.e., indirect climate effects and recovery from past land use) in driving understory responses at landscape scales? Stacked individual species distribution models fit with climate, forest, and soil predictors (248 species currently present in the landscape, derived from 150 field plots stratified by elevation and forest development, overall area under the receiving operator characteristic curve = 0.86) were driven with projected climate (RCP4.5 and RCP8.5) and modeled forest variables to predict plant community change. Nearly all species persisted in the landscape in 2050, but on average 8% of the species pool was lost by the end of the century. By 2100, landscape mean species richness and understory cover declined (-13% and -8%, respectively), warm-adapted species increasingly dominated plant communities (i.e., thermophilization, +12%), and plot-level turnover was high (62%). Subalpine forests experienced the greatest richness declines (-16%), most thermophilization (+17%), and highest turnover (67%), resulting in plant community homogenization across elevation zones. Climate rather than forest change was the dominant driver of understory responses. The magnitude of unabated 21st-century change is likely to erode plant diversity in a species richness hotspot, calling for stronger conservation and climate mitigation efforts.

Pollinator type strongly impacts gene flow within and among plant populations for six Neotropical species.

Animal pollinators directly affect plant gene flow by transferring pollen grains between individuals. Pollinators with restricted mobility are predicted to limit gene flow within and among populations, whereas pollinators that fly longer distances are likely to promote genetic cohesion. These predictions, however, remain poorly tested. We examined population genetic structure and fine-scale spatial genetic structure (FSGS) in six perennial understory angiosperms in Andean cloud forests of northwestern Ecuador. Species belong to three families (Gesneriaceae, Melastomataceae, and Rubiaceae), and within each family we paired one insect-pollinated with one hummingbird-pollinated species, predicting that insect-pollinated species have greater population differentiation (as quantified with the FST statistic) and stronger FSGS (as quantified with the SP statistic) than hummingbird-pollinated species. We confirmed putative pollinators through a literature review and fieldwork, and inferred population genetic parameters with a genome-wide genotyping approach. In two of the three species pairs, insect-pollinated species had much greater (>2-fold) population-level genetic differentiation and correspondingly steeper declines in fine-scale genetic relatedness. In the Gesneriaceae pair, however, FST and SP values were similar between species and to those of the other hummingbird-pollinated plants. In this pair, the insect pollinators are euglossine bees (as opposed to small bees and flies in the other pairs), which are thought to forage over large areas, and therefore may provide similar levels of gene flow as hummingbirds. Overall, our results shed light on how different animal pollination modes influence the spatial scale of plant gene flow, suggesting that small insects strongly decrease genetic cohesion.

Modeled Vegetation Community Trajectories: Effects from Climate Change, Atmospheric Nitrogen Deposition, and Soil Acidification Recovery.

Forest understory plant communities in the United States harbor most of the vegetation diversity of forests and are often sensitive to changes in climate and atmospheric deposition of nitrogen (N). As temperature increases from human-caused climate change and soils recover from long term atmospheric deposition of N and sulfur (S), it is unclear how these important ecosystem components will respond. We used the newly developed US-PROPS model - based on species response functions for over 1,500 species - to evaluate the potential impacts of atmospheric N deposition and climate change on species occurrence probability for a case study in the forested ecosystems of the Great Smoky Mountains National Park (GRSM), an iconic park in the southeastern United States. We evaluated six future scenarios from various combinations of two potential recoveries of soil pH (no change, +0.5 pH units) and three climate futures (no change, +1.5, +3.0 deg C). Species critical loads (CLs) of N deposition and projected responses for each scenario were determined. Critical loads were estimated to be low (< 2 kg N/ha/yr) to protect all species under current and expected future conditions across broad regions of GRSM and these CLs were exceeded at large spatial extents among scenarios. Northern hardwood, yellow pine, and chestnut oak forests were among the most N-sensitive vegetation map classes found within GRSM. Potential future air temperature conditions generally led to decreases in the maximum occurrence probability for species. Therefore, CLs were considered "unattainable" in these situations because the specified level of protection used for CL determination (i.e., maximum occurrence probability under ambient conditions) was not attainable. Although some species showed decreases in maximum occurrence probability with simulated increases in soil pH, most species were favored by increased pH. The importance of our study is rooted in the methodology described here for establishing regional CLs and for evaluating future conditions, which is transferable to other national parks in the U.S. and in Europe where the original PROPS model was developed.

Novelties from the herbaceous stratum in a key region for the conservation of the Southern Amazon / Novidades do estrato herbáceo em uma região chave para a conservação da Amazônia Meridional

Abstract The contribution of the herbaceous stratum to tropical plant diversity is considerable, however this component remains undersampled. We investigated floristic, structural, ecological and conservation issues concerning the herbaceous component of a seasonal deciduous forest associated with granitic rock outcrops in the Cristalino Region, a key area for biodiversity conservation in the Brazilian Amazon. We installed a permanent plot of 1 ha, allocating 10 transect-lines of 20 m each. We identified the sampled individuals, measured height and projection, and verified cover and frequency per species, genera and family. We recorded 86 species, 62 genera and 25 families, with Orchidaceae being the family with the highest species richness. Among the 26 new species added to Cristalino Flora, we included Philodendron deflexum Poepp. ex Schott and Griffinia nocturna Ravenna, the latter 'Critically Endangered'. Furthermore, the occurrence of G. nocturna in an Amazonian forest matrix is a novelty in this study. The estimate of species diversity according to Shannon-Wiener (H') was 2.43 nats.ind.-1 (equivalent to 11.37 ± 0.90 IC95% equally common species), and according to Simpson (1/D), 6.82 (± 0.648 IC95%). The rarefaction and extrapolation curves for the diversity estimates tended to stabilize. Although the vegetation on rock outcrops usually presents a high number of endemic species, this pattern was not found in our study area, which can be explained by its continuous occurrence in the forest matrix. The understory of our study area consists in a mixture of floras, being composed mainly of species from the Amazon and/or Cerrado biomes. In view of the current anthropic pressure faced by the southern Amazon, we reinforce the importance of carrying out inventories of its herbaceous communities, since the risk of species loss is even more alarming when considering present undersampling of this component.

Resumo A contribuição do estrato herbáceo para a diversidade de plantas tropicais é considerável, mas esse componente permanece subamostrado. Investigamos questões florísticas, estruturais, ecológicas e de conservação relacionadas ao componente herbáceo de uma floresta estacional decidual associada a afloramentos rochosos graníticos na região do Cristalino, que é uma área chave para a conservação da biodiversidade na Amazônia brasileira. Instalamos um plot permanente de 1 ha, alocando 10 linhas de 20 m cada. Identificamos os indivíduos amostrados, medimos altura e projeção e verificamos cobertura e frequência por espécie, gênero e família. Registramos 86 espécies, 62 gêneros e 25 famílias, sendo Orchidaceae a família com maior riqueza de espécies. Entre as 26 novas espécies adicionadas à Flora do Cristalino, incluímos Philodendron deflexum Poepp ex Schott e Griffinia nocturna Ravenna, esta última 'Criticamente Ameaçada'. Além disso, a ocorrência de G. nocturna em uma matriz florestal amazônica é uma novidade neste estudo. A estimativa da diversidade de espécies de acordo com Shannon-Wiener (H ') foi 2,43 nats / ind.-1 (equivalente a 11,37 ± 0,90 IC95% espécies igualmente comuns), e de acordo com Simpson (1/D), 6,82 (± 0,648 IC95%). As curvas de rarefação e extrapolação para as estimativas de diversidade tenderam à estabilização. Embora a vegetação associada a afloramentos rochosos em geral apresente um elevado número de espécies endêmicas, esse padrão não foi encontrado para a nossa área de estudo, o que pode ser explicado pela sua ocorrência contínua à matriz florestal. O sub-bosque da nossa área de estudo apresenta mistura de floras, sendo composto principalmente por espécies dos biomas Amazônia e/ou Cerrado. Diante das pressões antrópicas existentes no sul da Amazônia, reforçamos a importância da realização de inventários das comunidades herbáceas desse bioma, já que o risco de perda de espécies é ainda mais alarmante quando consideramos a subamostragem desse componente.

Shade Avoidance and Light Foraging of a Clonal Woody Species, Pachysandra terminalis.

(1) Background: A central subject in clonal plant ecology is to elucidate the mechanism by which clones forage resources in heterogeneous environments. Compared with studies conducted in laboratories or experimental gardens, studies on light foraging of forest woody clonal plants in their natural habitats are limited. (2) Methods: We investigated wild populations of an evergreen clonal understory shrub, Japanese pachysandra (Pachysandra terminalis Siebold & Zucc.), in two cool-temperate forests in Japan. (3) Results: Similar to the results of herbaceous clonal species, this species formed a dense stand in a relatively well-lit place, and a sparse stand in a shaded place. Higher specific rhizome length (i.e., length per unit mass) in shade resulted in lower ramet population density in shade. The individual leaf area, whole-ramet leaf area, or ramet height did not increase with increased light availability. The number of flower buds per flowering ramet increased as the canopy openness or population density increased. (4) Conclusions: Our results provide the first empirical evidence of shade avoidance and light foraging with morphological plasticity for a clonal woody species.

Threshold effects of air pollution and climate change on understory plant communities at forested sites in the eastern United States.

Forest understory plant communities in the eastern United States are often diverse and are potentially sensitive to changes in climate and atmospheric inputs of nitrogen caused by air pollution. In recent years, empirical and processed-based mathematical models have been developed to investigate such changes in plant communities. In the study reported here, a robust set of understory vegetation response functions (expressed as version 2 of the Probability of Occurrence of Plant Species model for the United States [US-PROPS v2]) was developed based on observations of forest understory and grassland plant species presence/absence and associated abiotic characteristics derived from spatial datasets. Improvements to the US-PROPS model, relative to version 1, were mostly focused on inclusion of additional input data, development of custom species-level input datasets, and implementation of methods to address uncertainty. We investigated the application of US-PROPS v2 to evaluate the potential impacts of atmospheric nitrogen (N) and sulfur (S) deposition, and climate change on forest ecosystems at three forested sites located in New Hampshire, Virginia, and Tennessee in the eastern United States. Species-level N and S critical loads (CLs) were determined under ambient deposition at all three modeled sites. The lowest species-level CLs of N deposition at each site were between 2 and 11 kg N/ha/yr. Similarly, the lowest CLs of S deposition, based on the predicted soil pH response, were less than 2 kg S/ha/yr among the three sites. Critical load exceedance was found at all three model sites. The New Hampshire site included the largest percentage of species in exceedance. Simulated warming air temperature typically resulted in lower maximum occurrence probability, which contributed to lower CLs of N and S deposition. The US-PROPS v2 model, together with the PROPS-CLF model to derive CL functions, can be used to develop site-specific CLs for understory plants within broad regions of the United States. This study demonstrates that species-level CLs of N and S deposition are spatially variable according to the climate, light availability, and soil characteristics at a given location. Although the species niche models generally performed well in predicting occurrence probability, there remains uncertainty with respect to the accuracy of reported CLs. As such, the specific CLs reported here should be considered as preliminary estimates.

Effects of earthworms and white-tailed deer on roots, arbuscular mycorrhizae, and forest seedling performance.

Changes in understory plant composition and biodiversity declines in northeastern North American forests are widespread. Preserving species and ecosystem function requires appropriate identification and management of important stressors. Coexistence of stressors, among them earthworm invasions and white-tailed deer, makes correct identification of mechanisms that cause diversity declines challenging. We used an established factorial experiment to assess survival and growth of native seedlings (Actaea pachypoda, Aquilegia canadensis, Cornus racemosa, Quercus rubra, and Prenanthes alba) in response to presence/absence of deer and earthworms. We expected deer and earthworms to reduce seedling survival and biomass, and we evaluated potential pathways to explain this impact (soil N and P concentrations and pools, root architecture, and arbuscular mycorrhizal fungi [AMF] colonization). We developed structural equation models (SEM) to identify specific pathways through which earthworms and deer were impacting plant species with different life histories. Seedling survival was not affected by our treatments nor the plant and soil variables we tested. Actaea biomass was smaller in earthworm-invaded plots, and with larger total N pools. In contrast, both deer and earthworm treatments were associated with lower soil nutrient concentrations, and earthworm-invaded plots had smaller N and extractable P pools. Actaea, Cornus, Prenanthes, and Quercus seedlings had a lower proportion of fine roots in earthworm-invaded plots, while fine roots in Aquilegia made up a higher proportion of the root system. AMF colonization in Quercus was reduced in sites colonized by earthworms, but AMF in other species were unaffected. Our SEMs showed high correlation among soil variables, but because we do not know which variables are drivers of this change and which are passengers, we can only conclude that they are changing together as deer and earthworms exert their respective influence. Different plant species responded in idiosyncratic ways to earthworm and deer effects on soil fertility, root architecture and limited effects on AMF colonization. While earthworm and deer-mediated changes to fine roots, soil nutrients, and AMF may lead to changes in plant performance over time, these changes rarely translated to lower plant performance in our seedlings.

Early spring warming may hasten leaf emergence in Erythronium americanum.

PREMISE: Climate change is making spring arrive earlier than in the past, causing some species to alter the timing of their spring activities. This study addressed whether Erythronium americanum Ker Gawl. (trout lily), a common spring ephemeral, can emerge earlier if exposed to early spring warming. METHODS: I collected corms of Erythronium americanum in the fall, overwintered them in soil, and exposed them to warming in either mid (early treatment) or late (late treatment) February. The timing of leaf emergence was monitored and compared between treatments. RESULTS: Leaves exposed to early warming emerged earlier than those in the late treatment. Bud break happened closer to date of exposure to warming in the late treatment than in the early treatment. CONCLUSIONS: Spring ephemerals may be able to produce leaves early in response to early spring warming induced by climate change. Risk of late frost and eventual shading by the canopy may limit the duration of a potentially extended growing season.

Utilizing herbarium specimens to quantify historical mycorrhizal communities.

PREMISE OF THE STUDY: Mycorrhiza are critical to ecosystem functioning, but a lack of historical baseline data limits our understanding of the long-term belowground effects of global change. Herbarium specimens may provide this needed insight. However, it is unknown whether DNA of arbuscular mycorrhizal fungi (AMF) can be reliably extracted from vascular plant specimen roots. METHODS: We sampled roots from herbarium specimens of four herbaceous forest species collected in western Pennsylvania between 1881-2008. Using molecular methods (terminal restriction fragment length polymorphism and sequence analysis), we quantified AMF communities from specimen roots and tested for contamination. RESULTS: We successfully amplified AMF DNA from 44% (21/48) of the root but not leaf samples, indicating specimen contamination was negligible. As expected, there were significant differences in AMF composition between plant species (P < 0.05). However, no differences in AMF communities were detected through time, possibly due to limited sample size and low amplification rates in recent collections. DISCUSSION: Herbaria have potential as sources of valuable belowground microbial data to answer questions across geographic, temporal, and taxonomic scales otherwise not feasible. Ongoing methodological developments will only magnify this potential. Further tests are needed to determine curatorial practices that maximize this innovative use of herbarium specimens.

Carbon gain phenologies of spring-flowering perennials in a deciduous forest indicate a novel niche for a widespread invader.

Strategies of herbaceous species in deciduous forests are often characterized by the timing of life history phases (e.g. emergence, flowering, leaf senescence) relative to overstory tree canopy closure. Although springtime photosynthesis is assumed to account for the majority of their annual carbon budgets, the 12-month photosynthetic trajectories of forest herbs have not been quantified. We measured the temporal dynamics of carbon assimilation for seven native herbaceous perennials and the biennial Alliaria petiolata, a widespread invader in eastern North American forests. We assessed the relative importance of spring, summer, and autumn to species-level annual carbon budgets. Spring-emerging species showed significant variation in carbon assimilation patterns. High spring irradiance before canopy closure accounted for 39-100% of species-level annual carbon assimilation, but summer and autumn accounted for large proportions of some species' carbon budgets (up to 58% and 19%, respectively). Alliaria was phenologically unique, taking advantage both autumn and spring irradiance. Although spring-emerging understory species are often expected to rely on early-season irradiance, our results highlight interspecific differences and the importance of mid-late season carbon gain. Phenological strategies of forest herbs are a continuum rather than discrete categories, and invasive species may follow strategies that are underrepresented in the native flora.

Chronic impacts of invasive herbivores on a foundational forest species: a whole-tree perspective.

Forests make up a large portion of terrestrial plant biomass, and the long-lived woody plants that dominate them possess an array of traits that deter consumption by forest pests. Although often extremely effective against native consumers, invasive species that avoid or overcome these defenses can wreak havoc on trees and surrounding ecosystems. This is especially true when multiple invasive species co-occur, since interactions between invasive herbivores may yield non-additive effects on the host. While the threat posed by invasive forest pests is well known, long-term field experiments are necessary to explore these consumer-host interactions at appropriate spatial and temporal scales. Moreover, it is important to measure multiple variables to get a "whole-plant" picture of their combined impact. We report the results of a 4-yr field experiment addressing the individual and combined impacts of two invasive herbivores, the hemlock woolly adelgid (Adelges tsugae) and elongate hemlock scale (Fiorinia externa), on native eastern hemlock (Tsuga canadensis) in southern New England. In 2011, we planted 200 hemlock saplings into a temperate forest understory and experimentally manipulated the presence/absence of both herbivore species; in 2015, we harvested the 88 remaining saplings and assessed plant physiology, growth, and resource allocation. Adelgids strongly affected hemlock growth: infested saplings had lower above/belowground biomass ratios, more needle loss, and produced fewer new needles than control saplings. Hemlock scale did not alter plant biomass allocation or growth, and its co-occurrence did not alter the impact of adelgid. While both adelgid and scale impacted the concentrations of primary metabolites, adelgid effects were more pronounced. Adelgid feeding simultaneously increased free amino acids local to feeding sites and a ~30% reduction in starch. The cumulative impact of adelgid-induced needle loss, manipulation of nitrogen pools, and the loss of stored resources likely accelerates host decline through disruption of homeostatic source-sink dynamics occurring at the whole-plant level. Our research stresses the importance of considering long-term impacts to predict how plants will cope with contemporary pressures experienced in disturbed forests.

Vegetation dynamics associated with changes in atmospheric nitrogen deposition and climate in hardwood forests of Shenandoah and Great Smoky Mountains National Parks, USA.

Ecological effects of atmospheric nitrogen (N) and sulfur (S) deposition on two hardwood forest sites in the eastern United States were simulated in the context of a changing climate using the dynamic coupled biogeochemical/ecological model chain ForSAFE-Veg. The sites are a mixed oak forest in Shenandoah National Park, Virginia (Piney River) and a mixed oak-sugar maple forest in Great Smoky Mountains National Park, Tennessee (Cosby Creek). The sites have received relatively high levels of both S and N deposition and the climate has warmed over the past half century or longer. The model was used to evaluate the composition of the understory plant communities, the alignment between plant species niche preferences and ambient conditions, and estimate changes in relative species abundances as reflected by plant cover under various scenarios of future atmospheric N and S deposition and climate change. The main driver of ecological effects was soil solution N concentration. Results of this research suggested that future climate change might compromise the capacity for the forests to sustain habitat suitability. However, vegetation results should be considered preliminary until further model validation can be performed. With expected future climate change, preliminary estimates suggest that sustained future N deposition above 7.4 and 5.0 kg N/ha/yr is expected to decrease contemporary habitat suitability for indicator plant species located at Piney River and Cosby Creek, respectively.

Feasibility of coupled empirical and dynamic modeling to assess climate change and air pollution impacts on temperate forest vegetation of the eastern United States.

Changes in climate and atmospheric nitrogen (N) deposition caused pronounced changes in soil conditions and habitat suitability for many plant species over the latter half of the previous century. Such changes are expected to continue in the future with anticipated further changing air temperature and precipitation that will likely influence the effects of N deposition. To investigate the potential long-term impacts of atmospheric N deposition on hardwood forest ecosystems in the eastern United States in the context of climate change, application of the coupled biogeochemical and vegetation community model VSD+PROPS was explored at three sites in New Hampshire, Virginia, and Tennessee. This represents the first application of VSD+PROPS to forest ecosystems in the United States. Climate change and elevated (above mid-19th century) N deposition were simulated to be important factors for determining habitat suitability. Although simulation results suggested that the suitability of these forests to support the continued presence of their characteristic understory plant species might decline by the year 2100, low data availability for building vegetation response models with PROPS resulted in uncertain results at the extremes of simulated N deposition. Future PROPS model development in the United States should focus on inclusion of additional foundational data or alternate candidate predictor variables to reduce these uncertainties.

Assessing plant community composition fails to capture impacts of white-tailed deer on native and invasive plant species.

Excessive herbivory can have transformative effects on forest understory vegetation, converting diverse communities into depauperate ones, often with increased abundance of non-native plants. White-tailed deer are a problematic herbivore throughout much of eastern North America and alter forest understory community structure. Reducing (by culling) or eliminating (by fencing) deer herbivory is expected to return understory vegetation to a previously diverse condition. We examined this assumption from 1992 to 2006 at Fermilab (Batavia, IL) where a cull reduced white-tailed deer (Odocoileus virginianus) abundance in 1998/1999 by 90 % from 24.6 to 2.5/km2, and at West Point, NY, where we assessed interactive effects of deer, earthworms, and invasive plants using 30 × 30 m paired fenced and open plots in 12 different forests from 2009 to 2012. We recorded not only plant community responses (species presence and cover) within 1 m2 quadrats, but also responses of select individual species (growth, reproduction). At Fermilab, introduced Alliaria petiolata abundance initially increased as deer density increased, but then declined after deer reduction. The understory community responded to the deer cull by increased cover, species richness and height, and community composition changed but was dominated by early successional native forbs. At West Point plant community composition was affected by introduced earthworm density but not deer exclusion. Native plant cover increased and non-native plant cover decreased in fenced plots, thus keeping overall plant cover similar. At both sites native forb cover increased in response to deer reduction, but the anticipated response of understory vegetation failed to materialize at the community level. Deer-favoured forbs (Eurybia divaricata, Maianthemum racemosum, Polygonatum pubescens and Trillium recurvatum) grew taller and flowering probability increased in the absence of deer. Plant community monitoring fails to capture initial and subtle effects of reduced or even cessation of deer browse on browse sensitive species. Measuring responses of individual plants (growth, flowering and reproductive success) provides a more sensitive and powerful assessment of forest understory responses to deer management.

Tracking lags in historical plant species' shifts in relation to regional climate change.

Can species shift their distributions fast enough to track changes in climate? We used abundance data from the 1950s and the 2000s in Wisconsin to measure shifts in the distribution and abundance of 78 forest-understory plant species over the last half-century and compare these shifts to changes in climate. We estimated temporal shifts in the geographic distribution of each species using vectors to connect abundance-weighted centroids from the 1950s and 2000s. These shifts in distribution reflect colonization, extirpation, and changes in abundance within sites, separately quantified here. We then applied climate analog analyses to compute vectors representing the climate change that each species experienced. Species shifted mostly to the northwest (mean: 49 ± 29 km) primarily reflecting processes of colonization and changes in local abundance. Analog climates for these species shifted even further to the northwest, however, exceeding species' shifts by an average of 90 ± 40 km. Most species thus failed to match recent rates of climate change. These lags decline in species that have colonized more sites and those with broader site occupancy, larger seed mass, and higher habitat fidelity. Thus, species' traits appear to affect their responses to climate change, but relationships are weak. As climate change accelerates, these lags will likely increase, potentially threatening the persistence of species lacking the capacity to disperse to new sites or locally adapt. However, species with greater lags have not yet declined more in abundance. The extent of these threats will likely depend on how other drivers of ecological change and interactions among species affect their responses to climate change.

Heteroploid Knautia drymeia includes K. gussonei and cannot be separated into diagnosable subspecies.

PREMISE OF THE STUDY: Knautia drymeia is a morphologically variable, diploid and tetraploid temperate forest understory species distributed in southeastern Europe and adjacent areas. The species is an excellent system to explore the influence of polypoidy on taxonomic delineations, the role of hybridization among genetically distant populations in polyploid evolution, and the impact of glacial refugia on the evolution of polyploids. METHODS: Amplified fragment length polymorphism fingerprinting and multivariate analyses of morphological characters were performed on 57 populations spanning the distribution area of K. drymeia. K-means clustering, comparison of in-silico tetraploids and observed tetraploids, and a phylogeographic analysis using relaxed random walks were used to explore the genetic structure within the diploids, to infer the origin of the tetraploids and to reconstruct range expansion through time. Further, we contrasted the morphology and genetic groups with current taxonomy and evaluated the status of the tetraploid Apennine endemic K. gussonei and the intraspecific taxa of K. drymeia. KEY RESULTS: The genetic structure was strongly geographically correlated and yielded four genetic groups; K. gussonei was inseparable from K. drymeia. Distributions of diploid lineages are suggestive of glacial refugia in the northwesternmost and southeastern Balkan Peninsula. Polyploids originated at least two times, as autopolyploids and probably additionally also as allopolyploids. Morphological divergence corresponded with neither genetic groups nor current taxonomy. CONCLUSIONS: Genetic and morphometric data confirmed neither divergence of K. gussonei nor recognition of subspecies within K. drymeia. We therefore propose treating K. drymeia as a morphologically and genetically variable species without infraspecific taxa.

Forest understory plant and soil microbial response to an experimentally induced drought and heat-pulse event: the importance of maintaining the continuum.

Drought duration and intensity are expected to increase with global climate change. How changes in water availability and temperature affect the combined plant-soil-microorganism response remains uncertain. We excavated soil monoliths from a beech (Fagus sylvatica L.) forest, thus keeping the understory plant-microbe communities intact, imposed an extreme climate event, consisting of drought and/or a single heat-pulse event, and followed microbial community dynamics over a time period of 28 days. During the treatment, we labeled the canopy with (13) CO2 with the goal of (i) determining the strength of plant-microbe carbon linkages under control, drought, heat and heat-drought treatments and (ii) characterizing microbial groups that are tightly linked to the plant-soil carbon continuum based on (13) C-labeled PLFAs. Additionally, we used 16S rRNA sequencing of bacteria from the Ah horizon to determine the short-term changes in the active microbial community. The treatments did not sever within-plant transport over the experiment, and carbon sinks belowground were still active. Based on the relative distribution of labeled carbon to roots and microbial PLFAs, we determined that soil microbes appear to have a stronger carbon sink strength during environmental stress. High-throughput sequencing of the 16S rRNA revealed multiple trajectories in microbial community shifts within the different treatments. Heat in combination with drought had a clear negative effect on microbial diversity and resulted in a distinct shift in the microbial community structure that also corresponded to the lowest level of label found in the PLFAs. Hence, the strongest changes in microbial abundances occurred in the heat-drought treatment where plants were most severely affected. Our study suggests that many of the shifts in the microbial communities that we might expect from extreme environmental stress will result from the plant-soil-microbial dynamics rather than from direct effects of drought and heat on soil microbes alone.

Light acclimation of photosynthesis in two closely related firs (Abies pinsapo Boiss. and Abies alba Mill.): the role of leaf anatomy and mesophyll conductance to CO2.

Leaves growing in the forest understory usually present a decreased mesophyll conductance (gm) and photosynthetic capacity. The role of leaf anatomy in determining the variability in gm among species is known, but there is a lack of information on how the acclimation of gm to shade conditions is driven by changes in leaf anatomy. Within this context, we demonstrated that Abies pinsapo Boiss. experienced profound modifications in needle anatomy to drastic changes in light availability that ultimately led to differential photosynthetic performance between trees grown in the open field and in the forest understory. In contrast to A. pinsapo, its congeneric Abies alba Mill. did not show differences either in needle anatomy or in photosynthetic parameters between trees grown in the open field and in the forest understory. The increased gm values found in trees of A. pinsapo grown in the open field can be explained by occurrence of stomata at both needle sides (amphistomatous needles), increased chloroplast surface area exposed to intercellular airspace, decreased cell wall thickness and, especially, decreased chloroplast thickness. To the best of our knowledge, the role of such drastic changes in ultrastructural needle anatomy in explaining the response of gm to the light environment has not been demonstrated in field conditions.

A reference-based approach for estimating leaf area and cover in the forest herbaceous layer.

Cover data are used to assess vegetative response to a variety of ecological factors. Estimating cover in the herbaceous layer of forests presents a problem because the communities are structurally complex and rich in species. The currently employed techniques for estimating cover are less than optimal for measuring such rich understories because they are inaccurate, slow, or impracticable. A reference-based approach to estimating cover is presented that compares the area of foliar surfaces to the area of an observer's hand. While this technique has been used to estimate cover in prior studies, its accuracy has not been tested. We tested this hand-area method at the individual plant, population, and community scales in a deciduous forest herbaceous layer, and in a separate farm experiment. The precision, accuracy, observer bias, and species bias of the method were tested by comparing the hand-estimated leaf area index values with actual leaf area index, measured using a leaf area meter. The hand-area method was very precise when regressed against actual leaf area index at the plant, population, and community scales (R(2) of 0.97, 0.93, and 0.87). Among the deciduous sites, the hand-area method overestimated leaf area index consistently by 39.1% at all scales. There was no observer bias detected at any scale, but plant overestimation bias was detected in one species at the population scale. The hand-area method is a rapid and reliable technique for estimating leaf area index or cover in the forest herbaceous layer and should be useful to field ecologists interested in answering questions at the plant, population, or community level.