Linear relationships between aboveground biomass and plant cover in low open herbaceous vegetation

Abstract. Four herbaceous plant species of a sand dune area and several herb species of an open early‐successional patch were investigated for the occurrence of a simple relationship between aboveground biomass and plant cover. Without exception linear regressions of aboveground biomass on plant cover were found with slope factors depending on the growth form of the species. These results suggest that (early) growth of herbaceous plants in low and (temporarily) open vegetation is not affected by possible constraints caused by a decreasing ratio of plant cover to aboveground biomass. The obtained linear relationships could be used for rapid non‐destructive determination of aboveground biomass by image‐analysis of cover data.     

Nonintrusive Field Experiments Show Different Plant Responses to Warming and Drought Among Sites,Seasons, and Species in a North–South European Gradient

We used a novel, nonintrusive experimental system to examine plant responses to warming and drought across a climatic and geographical latitudinal gradient of shrubland ecosystems in four sites from northern to southern Europe (UK, Denmark, The Netherlands, and Spain). In the first two years of experimentation reported here, we measured plant cover and biomass by the pinpoint method, plant ¹⁴C uptake, stem and shoot growth, flowering, leaf chemical concentration, litterfall, and herbivory damage in the dominant plant species of each site. The two years of approximately 1°C experimental warming induced a 15% increase in total aboveground plant biomass growth in the UK site. Both direct and indirect effects of warming, such as longer growth season and increased nutrient availability, are likely to be particularly important in this and the other northern sites which tend to be temperature-limited. In the water-stressed southern site, there was no increase in total aboveground plant biomass growth as expected since warming increases water loss, and temperatures in those ecosystems are already close to the optimum for photosynthesis. The southern site presented instead the most negative response to the drought treatment consisting of a soil moisture reduction at the peak of the growing season ranging from 33% in the Spanish site to 82% in The Netherlands site. In the Spanish site there was a 14% decrease in total aboveground plant biomass growth relative to control. Flowering was decreased by drought (up to 24% in the UK and 40% in Spain). Warming and drought decreased litterfall in The Netherlands site (33% and 37%, respectively) but did not affect it in the Spanish site. The tissue P concentrations generally decreased and the N/P ratio increased with warming and drought except in the UK site, indicating a progressive importance of P limitation as a consequence of warming and drought. The magnitude of the response to warming and drought was thus very sensitive to differences among sites (cold-wet northern sites were more sensitive to warming and the warm-dry southern site was more sensitive to drought), seasons (plant processes were more sensitive to warming during the winter than during the summer), and species. As a result of these multiple plant responses, ecosystem and community level consequences may be expected.     

The response of Alaskan arctic tundra to experimental warming: differences between short- and long-term responses

Global climate models predict continued rapid warming for most of the Arctic throughout the next century. To further understand the response of arctic tundra to climate warming, four sites in northern Alaska were warmed for five to seven consecutive growing seasons using open‐top chambers. Sites were located in dry heath and wet meadow communities near Barrow (71°18′N, 156°40′W) and Atqasuk (70°29′N, 157°25′W). Change in plant community composition was measured using a point frame method. During the period of observation, species richness declined in control plots by up to 2.7 species plot^?1. Responses to warming varied by site but similar trends included increased canopy height (?0.1 to 2.3 cm) and relative cover of standing dead plant matter (1.5–6.0%) and graminoids (1.8–5.8%) and decreased species diversity (0.1–1.7 species plot^?1) and relative cover of lichens (0.2–9.1%) and bryophytes (1.4–4.6%) (parentheses enclose the range of average values for the sites). The response to warming was separated into an initial short‐term response assessed after two growing seasons of warming and a secondary longer‐term response assessed after an additional three to five growing seasons of warming. The initial responses to warming were similar in the four sites, while the secondary responses varied by site. The response to warming was greater at Barrow than Atqasuk because of a greater initial response at Barrow. However, the long‐term response to warming was projected to be greater at Atqasuk because of a greater secondary response at Atqasuk. These findings show that predictions of vegetation change due to climate warming based on manipulative experiments will differ depending on both the duration and plant community on which the study focuses.     

Additive Effects of Warming and Increased Nitrogen Deposition in a Temperate Old Field: Plant Productivity and the Importance of Winter

Both climate warming and atmospheric nitrogen (N) deposition are predicted to alter plant productivity and species composition over the next century. However, the extent to which their effects may interact is unclear. For example, over winter, the effects of warming on soil freezing dynamics may promote ecosystem N losses, thereby limiting increases in productivity in response to warming, yet these losses may be compensated for by increased N deposition. We measured plant production and species composition in response to warming (winter-only or year-round) and N addition in a temperate old field. We used shoot allometric relationships to estimate aboveground production non-destructively and sampled root biomass destructively throughout two growing seasons. We also used spectral data (normalized difference vegetation index—NDVI) to examine the treatment effects on the timing of plant green-up and senescence. In 2007, which featured an exceptionally dry summer, there were no treatment effects on plant growth. However, in 2008, warming (both winter-only and year-round) and N addition combined approximately doubled aboveground productivity, and these effects were additive. Warming increased root biomass, but no N effect was evident. Conversely, N addition increased NDVI, but NDVI was unresponsive to warming. Overall, our results do not support the hypothesis that warming-induced changes to soil freezing dynamics limit plant productivity in our system. On the contrary, they demonstrate that winter warming alone can increase primary productivity to the same extent as year-round warming, and that this effect may interact very strongly with interannual variation in precipitation.     

Circumpolar arctic tundra biomass and productivity dynamics in response to projected climate change and herbivory

Satellite remote sensing data have indicated a general ‘greening’ trend in the arctic tundra biome. However, the observed changes based on remote sensing are the result of multiple environmental drivers, and the effects of individual controls such as warming, herbivory, and other disturbances on changes in vegetation biomass, community structure, and ecosystem function remain unclear. We apply ArcVeg, an arctic tundra vegetation dynamics model, to estimate potential changes in vegetation biomass and net primary production (NPP) at the plant community and functional type levels. ArcVeg is driven by soil nitrogen output from the Terrestrial Ecosystem Model, existing densities of Rangifer populations, and projected summer temperature changes by the NCAR CCSM4.0 general circulation model across the Arctic. We quantified the changes in aboveground biomass and NPP resulting from (i) observed herbivory only; (ii) projected climate change only; and (iii) coupled effects of projected climate change and herbivory. We evaluated model outputs of the absolute and relative differences in biomass and NPP by country, bioclimate subzone, and floristic province. Estimated potential biomass increases resulting from temperature increase only are approximately 5% greater than the biomass modeled due to coupled warming and herbivory. Such potential increases are greater in areas currently occupied by large or dense Rangifer herds such as the Nenets‐occupied regions in Russia (27% greater vegetation increase without herbivores). In addition, herbivory modulates shifts in plant community structure caused by warming. Plant functional types such as shrubs and mosses were affected to a greater degree than other functional types by either warming or herbivory or coupled effects of the two.     

Detecting changes in arctic tundra plant communities in response to warming over decadal time scales

Detecting the response of vegetation to climate forcing as distinct from spatial and temporal variability may be difficult, if not impossible, over the typical duration of most field studies. We analyzed the spatial and interannual variability of plant functional type biomass from field studies in low arctic tussock tundra and compared these to climate change simulations of plant community composition using a dynamic tundra vegetation model (ArcVeg). Spatial heterogeneity of peak season live aboveground biomass was estimated using field samples taken from low arctic tundra at Ivotuk, Alaska (68.5°N, 155.7°W) in 1999. Coefficients of variation for live aboveground biomass at the 1 m² scale ranged from 14.6% for deciduous shrubs, 18.5% for graminoids and 25.3% for mosses to over 57% for forbs and lichens. Spatial heterogeneity in the ArcVeg dynamic vegetation model was simulated to be greater than the field data, ranging from 37.1% for deciduous shrubs to 107.9% for forbs. Disturbances in the model, such as caribou grazing and freezing–thawing of soil, as well as demographic stochasticity, led to the greater variability in the simulated results. Temporal variances of aboveground live biomass over a 19-year period using data from Toolik Lake, AK fell within the range of field and simulation spatial variances. However, simulations using ArcVeg suggest that temporal variability can be substantially less than site-scale spatial variability. Field data coupled with ArcVeg simulations of climate change scenarios indicate that some changes in plant community composition may be detectable within two decades following the onset of warming, and shrubs and mosses might be the key indicators of community change. Model simulations also project increasing landscape scale spatial heterogeneity (particularly of shrubs) with increasing temperatures.     

Interactions of herbivore exclusion with warming and N addition in a grass-dominated temperate old field

Field experiments used to explore the effects of global change drivers, such as warming and nitrogen deposition on plant productivity and species composition, have typically focused on bottom-up processes. However, both direct and indirect responses of herbivores to the treatments could result in important interactions between top-down and bottom-up effects. These interactions may be complicated by the simultaneous effects of multiple herbivore taxa. We used rodent and mollusc exclosures in the plots of a warming and N addition field experiment to examine how herbivore removal would influence plant biomass responses to the treatments. The effect of rodent exclusion on grass biomass more than doubled in response to nitrogen addition, but did not respond to warming, whereas the effect of mollusc exclusion on grass biomass increased in response to warming, but not nitrogen. In contrast, the effect of rodent exclusion on total biomass (grasses and forbs combined) increased in response to both nitrogen and warming, while the effect of mollusc exclusion on total biomass was insensitive to nitrogen and warming. In no cases were there interactions between nitrogen and warming with respect to their influence on exclosure effects. Overall, our results demonstrated substantial and variable effects of multiple herbivore taxa on plant biomass responses to warming and N addition, despite the absence of conspicuous damage to the plant canopy. These results therefore highlight the potential importance of interactions between top-down and bottom-up factors in global change field experiments.     

Responses of Plant Community Composition and Biomass Production to Warming and Nitrogen Deposition in a Temperate Meadow Ecosystem

Climate change has profound influences on plant community composition and ecosystem functions. However, its effects on plant community composition and biomass production are not well understood. A four-year field experiment was conducted to examine the effects of warming, nitrogen (N) addition, and their interactions on plant community composition and biomass production in a temperate meadow ecosystem in northeast China. Experimental warming had no significant effect on plant species richness, evenness, and diversity, while N addition highly reduced the species richness and diversity. Warming tended to reduce the importance value of graminoid species but increased the value of forbs, while N addition had the opposite effect. Warming tended to increase the belowground biomass, but had an opposite tendency to decrease the aboveground biomass. The influences of warming on aboveground production were dependent upon precipitation. Experimental warming had little effect on aboveground biomass in the years with higher precipitation, but significantly suppressed aboveground biomass in dry years. Our results suggest that warming had indirect effects on plant production via its effect on the water availability. Nitrogen addition significantly increased above- and below-ground production, suggesting that N is one of the most important limiting factors determining plant productivity in the studied meadow steppe. Significant interactive effects of warming plus N addition on belowground biomass were also detected. Our observations revealed that environmental changes (warming and N deposition) play significant roles in regulating plant community composition and biomass production in temperate meadow steppe ecosystem in northeast China.     

Climatic and biotic extreme events moderate long‐term responses of above‐ and belowground sub‐Arctic heathland communities to climate change

Climate change impacts are not uniform across the Arctic region because interacting factors causes large variations in local ecosystem change. Extreme climatic events and population cycles of herbivores occur simultaneously against a background of gradual climate warming trends and can redirect ecosystem change along routes that are difficult to predict. Here, we present the results from sub‐Arctic heath vegetation and its belowground micro‐arthropod community in response to the two main drivers of vegetation damage in this region: extreme winter warming events and subsequent outbreaks of the defoliating autumnal moth caterpillar (Epirrita autumnata). Evergreen dwarf shrub biomass decreased (30%) following extreme winter warming events and again by moth caterpillar grazing. Deciduous shrubs that were previously exposed to an extreme winter warming event were not affected by the moth caterpillar grazing, while those that were not exposed to warming events (control plots) showed reduced (23%) biomass from grazing. Cryptogam cover increased irrespective of grazing or winter warming events. Micro‐arthropods declined (46%) following winter warming but did not respond to changes in plant community. Extreme winter warming and caterpillar grazing suppressed the CO₂ fluxes of the ecosystem. Evergreen dwarf shrubs are disadvantaged in a future sub‐Arctic with more stochastic climatic and biotic events. Given that summer warming may further benefit deciduous over evergreen shrubs, event and trend climate change may both act against evergreen shrubs and the ecosystem functions they provide. This is of particular concern given that Arctic heath vegetation is typically dominated by evergreen shrubs. Other components of the vegetation showed variable responses to abiotic and biotic events, and their interaction indicates that sub‐Arctic vegetation response to multiple pressures is not easy to predict from single‐factor responses. Therefore, while biotic and climatic events may have clear impacts, more work is needed to understand their net effect on Arctic ecosystems.     

Warming experiments elucidate the drivers of observed directional changes in tundra vegetation

Few studies have clearly linked long‐term monitoring with in situ experiments to clarify potential drivers of observed change at a given site. This is especially necessary when findings from a site are applied to a much broader geographic area. Here, we document vegetation change at Barrow and Atqasuk, Alaska, occurring naturally and due to experimental warming over nearly two decades. An examination of plant cover, canopy height, and community indices showed more significant differences between years than due to experimental warming. However, changes with warming were more consistent than changes between years and were cumulative in many cases. Most cases of directional change observed in the control plots over time corresponded with a directional change in response to experimental warming. These included increases in canopy height and decreases in lichen cover. Experimental warming resulted in additional increases in evergreen shrub cover and decreases in diversity and bryophyte cover. This study suggests that the directional changes occurring at the sites are primarily due to warming and indicates that further changes are likely in the next two decades if the regional warming trend continues. These findings provide an example of the utility of coupling in situ experiments with long‐term monitoring to accurately document vegetation change in response to global change and to identify the underlying mechanisms driving observed changes.     

Long-term ecosystem level experiments at Toolik Lake, Alaska, and at Abisko, Northern Sweden: generalizations and differences in ecosystem and plant type responses to global change

Long‐term ecosystem‐level experiments, in which the environment is manipulated in a controlled manner, are important tools to predict the responses of ecosystem functioning and composition to future global change. We present the results of a meta‐analysis performed on the results of long‐term ecosystem‐level experiments near Toolik Lake, Alaska, and Abisko, Sweden. We quantified aboveground biomass responses of different arctic and subarctic ecosystems to experimental fertilization, warming and shading. We not only analysed the general patterns but also the differences in responsiveness between sites and regions. Aboveground plant biomass showed a broad similarity of responses in both locations, and also showed some important differences. In both locations, aboveground plant biomass, particularly the biomass of deciduous and graminoid plants, responded most strongly to nutrient addition. The biomass of mosses and lichens decreased in both locations as the biomass of vascular plants increased. An important difference between the two regions was the smaller positive aboveground biomass response of deciduous shrubs in Abisko as compared with Toolik Lake. Whereas in Toolik Lake Betula nana increased its dominance and replaced many of the other plant types, in Abisko all vascular plant types increased in abundance without major shifts in relative abundance. The differences between the responses of the dominant vegetation types of the Toolik Lake region, i.e. tussock tundra systems, and that of the Abisko region, i.e. heath systems, may have important implications for ecosystem development under expected patterns of global change. However, there were also large site‐specific differences within each region. Several potential mechanistic explanations for the differences between sites and regions are discussed. The response patterns show the need for analyses of joint data sets from many regions and sites, in order to uncover common responses to changes in climate across large arctic regions from regional or local responses.     

Warming increases aboveground plant biomass and C stocks in vascular-plant-dominated Antarctic tundra

We passively warmed tundra on the Antarctic Peninsula over four growing seasons and assessed its effect on dry mass and C and N stocks associated with the vascular plants Colobanthus quitensis (a cushion‐forming forb) and Deschampsia antarctica (a tussock grass), and mosses. Temperature treatments involved a warmed treatment that raised diurnal and diel canopy air temperatures by 2.3 and 1.3 °C, respectively, and a near‐ambient temperature treatment that raised diurnal and diel temperatures by 0.2 °C. These two different temperature regimes were achieved by wrapping filters around the frames to different extents and were nested within three UV treatments that filtered different solar UV wavebands. The experiment also included an ambient control treatment (unfiltered frames), and supplemental water and fertilizer treatments (applied to unfiltered frames). After four growing seasons, we collected cores of each vascular plant species and assessed the mass and C and N content of the aboveground current‐year biomass, the litter layer (which included nongreen live stems), and the organic soil horizon (which included roots). The thin nature of the organic soil horizon allowed us to sample this complete horizon and estimate near‐total ecosystem C and N stocks. A comparison of the warmed and near‐ambient temperature treatments found that warming led to greater aboveground biomass of C. quitensis, and more C in the aboveground biomass of both vascular plant species. Warming resulted in lower N concentrations of the aboveground biomass of both species. The water use efficiency of both species was greater under warming, based on their higher δ¹³C values. The mass of the litter layer under C. quitensis was greater under warming, and this layer contained more C and N and had a higher C : N ratio. The mass of the organic soil horizon under both species was greater under warming, and this horizon also contained more C and N. Warming also changed the species composition of the plant community – cover of C. quitensis increased while that of mosses declined. Warming resulted in the input of biomass into the system that had greater C : N ratios (and was likely more recalcitrant to decomposition) because (1) warming increased the C : N ratio of the biomass produced by both vascular plant species, (2) these inputs increased with warming because of greater biomass production, and (3) increases in C. quitensis cover led to greater biomass inputs by this species and its biomass had a greater C : N ratio than D. antarctica. Water or fertilizer supplements had few effects on aboveground biomass or C and N concentrations or pools, consistent with the relatively wet maritime climate and high soil nutrient levels of this system. Total C pools in the aboveground biomass, litter, and organic soil horizon were greater under warming. Warmed plots contained from 272 to 319 g m⁻² more C than plots under near‐ambient temperatures, corresponding to a 23–34% increase in ecosystem C.     

Permafrost response to temperature rise in carbon and nutrient cycling: Effects from habitat‐specific conditions and factors of warming

Permafrost is experiencing climate warming at a rate that is two times faster than the rest of the Earth''s surface. However, it is still lack of a quantitative basis for predicting the functional stability of permafrost ecosystems in carbon (C) and nutrient cycling. We compiled the data of 708 observations from 89 air‐warming experiments in the Northern Hemisphere and characterized the general effects of temperature increase on permafrost C exchange and balance, biomass production, microbial biomass, soil nutrients, and vegetation N dynamics through a meta‐analysis. Also, an investigation was made on how responses might change with habitat‐specific (e.g., plant functional groups and soil moisture status) conditions and warming variables (e.g., warming phases, levels, and timing). The net ecosystem C exchange (NEE) was found to be downregulated by warming as a result of a stronger sensitivity to warming in respiration (15.6%) than in photosynthesis (6.2%). Vegetation usually responded to warming by investing more C to the belowground, as belowground biomass increased much more (30.1%) than aboveground biomass (2.9%). Warming had a minor effect on microbial biomass. Warming increased soil ammonium and nitrate concentrations. What''s more, a synthesis of 70 observations from 11 herbs and 9 shrubs revealed a 2.5% decline of N in green leaves. Compared with herbs, shrubs had a stronger response to respiration and had a decline in green leaf N to a greater extent. Not only in dry condition did green leaf N decline with warming but also in wet conditions. Warming in nongrowing seasons would negatively affect soil water, C uptake, and biomass production during growing seasons. Permafrost C loss and vegetation N decline may increase with warming levels and timing. Overall, these findings suggest that besides a positive C cycling–climate feedback, there will be a negative feedback between permafrost nutrient cycling and climate warming.     

Water availability drives aboveground biomass and bird richness in forest restoration plantings to achieve carbon and biodiversity cobenefits

To combat global warming and biodiversity loss, we require effective forest restoration that encourages recovery of species diversity and ecosystem function to deliver essential ecosystem services, such as biomass accumulation. Further, understanding how and where to undertake restoration to achieve carbon sequestration and biodiversity conservation would provide an opportunity to finance ecosystem restoration under carbon markets. We surveyed 30 native mixed‐species plantings in subtropical forests and woodlands in Australia and used structural equation modeling to determine vegetation, soil, and climate variables most likely driving aboveground biomass accrual and bird richness and investigate the relationships between plant diversity, aboveground biomass accrual, and bird diversity. We focussed on woodland and forest‐dependent birds, and functional groups at risk of decline (insectivorous, understorey‐nesting, and small‐bodied birds). We found that mean moisture availability strongly limits aboveground biomass accrual and bird richness in restoration plantings, indicating potential synergies in choosing sites for carbon and biodiversity purposes. Counter to theory, woody plant richness was a poor direct predictor of aboveground biomass accrual, but was indirectly related via significant, positive effects of stand density. We also found no direct relationship between aboveground biomass accrual and bird richness, likely because of the strong effects of moisture availability on both variables. Instead, moisture availability and patch size strongly and positively influenced the richness of woodland and forest‐dependent birds. For understorey‐nesting birds, however, shrub cover and patch size predicted richness. Stand age or area of native vegetation surrounding the patch did not influence bird richness. Our results suggest that in subtropical biomes, planting larger patches to higher densities, ideally using a diversity of trees and shrubs (characteristics of ecological plantings) in more mesic locations will enhance the provision of carbon and biodiversity cobenefits. Further, ecological plantings will aid the rapid recovery of woodland and forest bird richness, with comparable aboveground biomass accrual to less diverse forestry plantations.     

Response of plant species richness and primary productivity in shrublands along a north–south gradient in Europe to seven years of experimental warming and drought: reductions in primary productivity in the heat and drought year of 2003

We used a nonintrusive field experiment carried out at six sites – Wales (UK), Denmark (DK), the Netherlands (NL), Hungary (HU), Sardinia (Italy – IT), and Catalonia (Spain – SP) – along a climatic and latitudinal gradient to examine the response of plant species richness and primary productivity to warming and drought in shrubland ecosystems. The warming treatment raised the plot daily temperature by ca. 1 °C, while the drought treatment led to a reduction in soil moisture at the peak of the growing season that ranged from 26% at the SP site to 82% in the NL site. During the 7 years the experiment lasted (1999–2005), we used the pin‐point method to measure the species composition of plant communities and plant biomass, litterfall, and shoot growth of the dominant plant species at each site. A significantly lower increase in the number of species pin‐pointed per transect was found in the drought plots at the SP site, where the plant community was still in a process of recovering from a forest fire in 1994. No changes in species richness were found at the other sites, which were at a more mature and stable state of succession and, thus less liable to recruitment of new species. The relationship between annual biomass accumulation and temperature of the growing season was positive at the coldest site and negative at the warmest site. The warming treatment tended to increase the aboveground net primary productivity (ANPP) at the northern sites. The relationship between annual biomass accumulation and soil moisture during the growing season was not significant at the wettest sites, but was positive at the driest sites. The drought treatment tended to reduce the ANPP in the NL, HU, IT, and SP sites. The responses to warming were very strongly related to the Gaussen aridity index (stronger responses the lower the aridity), whereas the responses to drought were not. Changes in the annual aboveground biomass accumulation, litterfall, and, thus, the ANPP, mirrored the interannual variation in climate conditions: the most outstanding change was a decrease in biomass accumulation and an increase in litterfall at most sites during the abnormally hot year of 2003. Species richness also tended to decrease in 2003 at all sites except the cold and wet UK site. Species‐specific responses to warming were found in shoot growth: at the SP site, Globularia alypum was not affected, while the other dominant species, Erica multiflora, grew 30% more; at the UK site, Calluna vulgaris tended to grow more in the warming plots, while Empetrum nigrum tended to grow less. Drought treatment decreased plant growth in several studied species, although there were some species such as Pinus halepensis at the SP site or C. vulgaris at the UK site that were not affected. The magnitude of responses to warming and drought thus depended greatly on the differences between sites, years, and species and these multiple plant responses may be expected to have consequences at ecosystem and community level. Decreases in biodiversity and the increase in E. multiflora growth at the SP site as a response to warming challenge the assumption that sensitivity to warming may be less well developed at more southerly latitudes; likewise, the fact that one of the studied shrublands presented negative ANPP as a response to the 2003 heat wave also challenges the hypothesis that future climate warming will lead to an enhancement of plant growth and carbon sequestration in temperate ecosystems. Extreme events may thus change the general trend of increased productivity in response to warming in the colder sites.     

Non-Destructive Lichen Biomass Estimation in Northwestern Alaska: A Comparison of Methods

Terrestrial lichen biomass is an important indicator of forage availability for caribou in northern regions, and can indicate vegetation shifts due to climate change, air pollution or changes in vascular plant community structure. Techniques for estimating lichen biomass have traditionally required destructive harvesting that is painstaking and impractical, so we developed models to estimate biomass from relatively simple cover and height measurements. We measured cover and height of forage lichens (including single-taxon and multi-taxa “community” samples, n = 144) at 73 sites on the Seward Peninsula of northwestern Alaska, and harvested lichen biomass from the same plots. We assessed biomass-to-volume relationships using zero-intercept regressions, and compared differences among two non-destructive cover estimation methods (ocular vs. point count), among four landcover types in two ecoregions, and among single-taxon vs. multi-taxa samples. Additionally, we explored the feasibility of using lichen height (instead of volume) as a predictor of stand-level biomass. Although lichen taxa exhibited unique biomass and bulk density responses that varied significantly by growth form, we found that single-taxon sampling consistently under-estimated true biomass and was constrained by the need for taxonomic experts. We also found that the point count method provided little to no improvement over ocular methods, despite increased effort. Estimated biomass of lichen-dominated communities (mean lichen cover: 84.9±1.4%) using multi-taxa, ocular methods differed only nominally among landcover types within ecoregions (range: 822 to 1418 g m⁻²). Height alone was a poor predictor of lichen biomass and should always be weighted by cover abundance. We conclude that the multi-taxa (whole-community) approach, when paired with ocular estimates, is the most reasonable and practical method for estimating lichen biomass at landscape scales in northwest Alaska.     

Response of Alpine Plants to Nitrogen Addition on the Tibetan Plateau: A Meta-analysis

To identify the general effects of nitrogen addition on alpine plants, we used a meta-analysis approach to synthesize 599 observations from 51 studies on the Tibetan Plateau. Nitrogen addition significantly increased plant height by 19.0 %, plant biomass by 29.7 %, graminoid aboveground biomass by 89.8 %, and sedge aboveground biomass by 75.6 % but significantly decreased legume aboveground biomass by 34.5 %, forb aboveground biomass by 23.8 %, and species richness by 11.2 %. The effect of nitrogen addition on aboveground plant biomass and plant height increased with increasing the nitrogen addition rate. The effect of nitrogen addition on plant height increased with increasing mean annual precipitation but decreased with increasing mean annual temperature. Our findings suggested that the effect of nitrogen addition on alpine plants varied with plant functional types and nitrogen addition rate. In addition, climatic warming and precipitation changes may regulate the response of alpine plants to nitrogen addition on the Tibetan Plateau.     

Shifting grassland plant community structure drives positive interactive effects of warming and diversity on aboveground net primary productivity

Ecosystems worldwide are increasingly impacted by multiple drivers of environmental change, including climate warming and loss of biodiversity. We show, using a long‐term factorial experiment, that plant diversity loss alters the effects of warming on productivity. Aboveground primary productivity was increased by both high plant diversity and warming, and, in concert, warming (≈1.5 °C average above and belowground warming over the growing season) and diversity caused a greater than additive increase in aboveground productivity. The aboveground warming effects increased over time, particularly at higher levels of diversity, perhaps because of warming‐induced increases in legume and C4 bunch grass abundances, and facilitative feedbacks of these species on productivity. Moreover, higher plant diversity was associated with the amelioration of warming‐induced environmental conditions. This led to cooler temperatures, decreased vapor pressure deficit, and increased surface soil moisture in higher diversity communities. Root biomass (0–30 cm) was likewise consistently greater at higher plant diversity and was greater with warming in monocultures and at intermediate diversity, but at high diversity warming had no detectable effect. This may be because warming increased the abundance of legumes, which have lower root : shoot ratios than the other types of plants. In addition, legumes increase soil nitrogen (N) supply, which could make N less limiting to other species and potentially decrease their investment in roots. The negative warming × diversity interaction on root mass led to an overall negative interactive effect of these two global change factors on the sum of above and belowground biomass, and thus likely on total plant carbon stores. In total, plant diversity increased the effect of warming on aboveground net productivity and moderated the effect on root mass. These divergent effects suggest that warming and changes in plant diversity are likely to have both interactive and divergent impacts on various aspects of ecosystem functioning.     

Vegetation and soil responses to livestock grazing in Central Asian grasslands: a review of Chinese literature

Grasslands in northern China and the Qinghai-Tibetan plateau are particularly important to both ecosystem functioning and pastoral livelihoods. Although there are numerous degradation studies on the effect of livestock grazing across the region, they are largely only published in Chinese, and most focus on single sites. Based on case studies from 100 sites, covering a mean annual precipitation gradient of 95–744 mm, we present a comprehensive, internationally accessible review on the impact of livestock grazing on vegetation and soils. We compared ungrazed or slightly grazed sites with moderately and heavily grazed sites by evaluating changes in two indicator groups: vegetation (plant species richness, vegetation cover, aboveground biomass, belowground biomass and root/shoot ratio) and soil (pH, bulk density, organic C, total N, total P and available P). Most indicators declined with intensified grazing, while soil pH, bulk density and belowground biomass increased. Available P showed no clear response. Variables within indicator groups were mostly linearly correlated at a given grazing intensity. Relative grazing effects on different indicators varied along specific abiotic gradients. Grazing responses of plant species richness, aboveground biomass, soil bulk density, total N and available P interacted with precipitation patterns, while grazing effects on belowground biomass were influenced by temperature. Elevation had impact on grazing responses of aboveground biomass and soil organic carbon. Complex grazing effects reflect both methodological inconsistency and ecological complexity. Further assessments should consider specific characteristics of different indicators in the context of the local environment.     

Sawgrass (Cladium jamaicense) responses as early indicators of low-level phosphorus enrichment in the Florida Everglades

Anthropogenic phosphorus (P) inputs to the Florida Everglades have produced dramatic changes in the wetland vegetation of this otherwise oligotrophic system. While the proliferation of undesirable plant species in response to enrichment has been well documented, nutrient-related changes in the physiological and morphological attributes of existing vegetation, prior to any shifts in species composition or changes in the spatial extent of certain taxa, have yet to be adequately characterized. In this experiment, three sawgrass-dominated areas were enriched with P for 3 years at rates of 0.4 g P/m²/year (HP), 0.1 g P/m²/year (LP), or 0 g P/m²/year (controls) to assess potential impacts of P-enriched discharges from stormwater treatment areas into the Everglades. Elevated concentrations of TP in rhizomes and leaves and reduced ratios of leaf N:P were detected in HP plants within ~1 year at most sites. Live leaf densities, plant heights, and plant densities of the HP groups were generally higher than LP and control groups after 2 years, a pattern that was evident even after major fire events. Total aboveground biomass was significantly elevated in both HP and LP treatments at two of the three sites after 3 years. No change in species composition was detected during the study. Planned hydrologic restoration measures will increase P loads into parts of the Everglades that have not previously experienced anthropogenic P enrichment. Monitoring native vegetation such as sawgrass can be a sensitive and relatively robust means of detecting unintended P enrichment in these areas prior to shifts in vegetation community composition or changes in area cover of key species.