Variation in Detrital Resource Stoichiometry Signals Differential Carbon to Nutrient Limitation for Stream Consumers Across Biomes

Stoichiometric ratios of resources and consumers have been used to predict nutrient limitation across diverse terrestrial and aquatic ecosystems. In forested headwater streams, coarse and fine benthic organic matter (CBOM, FBOM) are primary basal resources for the food web, and the distribution and quality of these organic matter resources may therefore influence patterns of secondary production and nutrient cycling within stream networks or among biomes. We measured carbon (C), nitrogen (N), and phosphorus (P) content of CBOM and FBOM and calculated their stoichiometric ratios (C/N, C/P, N/P) from first- to fourth-order streams from tropical montane, temperate deciduous, and boreal forests, and tallgrass prairie, to compare the magnitude and variability of these resource types among biomes. We then used the ratios to predict nutritional limitations for consumers of each resource type. Across biomes, CBOM had consistently higher %C and %N, and higher and more variable C/N and C/P than FBOM, suggesting that microbial processing results in more tightly constrained elemental composition in FBOM than in CBOM. Biome-specific differences were observed in %P and N/P between the two resource pools; CBOM was lower in %P but higher in N/P than FBOM in the tropical montane and temperate deciduous forest biomes, while CBOM was higher in %P but similar in N/P than FBOM in the grassland and boreal forest biomes. Stable ¹³C isotopes suggest that FBOM likely derives from CBOM in tropical and temperate deciduous forest, but that additional non-detrital components may contribute to FBOM in boreal forests and grasslands. Comparisons of stoichiometric ratios of CBOM and FBOM to estimated needs of aquatic detritivores suggest that shredders feeding on CBOM are more likely to experience nutrient (N and/or P) than C limitation, whereas collector–gatherers consuming FBOM are more likely to experience C than N and/or P limitation. Our results suggest that differences in basal resource elemental content and stoichiometric ratios have the potential to affect consumer production and ecosystem rates of C, N, and P cycling in relatively consistent ways across diverse biomes.     

Maximum rooting depth of vegetation types at the global scale

The depth at which plants are able to grow roots has important implications for the whole ecosystem hydrological balance, as well as for carbon and nutrient cycling. Here we summarize what we know about the maximum rooting depth of species belonging to the major terrestrial biomes. We found 290 observations of maximum rooting depth in the literature which covered 253 woody and herbaceous species. Maximum rooting depth ranged from 0.3 m for some tundra species to 68 m for Boscia albitrunca in the central Kalahari; 194 species had roots at least 2 m deep, 50 species had roots at a depth of 5 m or more, and 22 species had roots as deep as 10 m or more. The average for the globe was 4.6±0.5 m. Maximum rooting depth by biome was 2.0±0.3 m for boreal forest. 2.1±0.2 m for cropland, 9.5±2.4 m for desert, 5.2±0.8 m for sclerophyllous shrubland and forest, 3.9±0.4 m for temperate coniferous forest, 2.9±0.2 m for temperate deciduous forest, 2.6±0.2 m for temperate grassland, 3.7±0.5 m for tropical deciduous forest, 7.3±2.8 m for tropical evergreen forest, 15.0±5.4 m for tropical grassland/savanna, and 0.5±0.1 m for tundra. Grouping all the species across biomes (except croplands) by three basic functional groups: trees, shrubs, and herbaceous plants, the maximum rooting depth was 7.0±1.2 m for trees, 5.1±0.8 m for shrubs, and 2.6±0.1 m for herbaceous plants. These data show that deep root habits are quite common in woody and herbaceous species across most of the terrestrial biomes, far deeper than the traditional view has held up to now. This finding has important implications for a better understanding of ecosystem function and its application in developing ecosystem models.     

Carbon balance in the tundra, boreal forest and humid tropical forest during climate change: scaling up from leaf physiology and soil carbon dynamics

Carbon exchange by the terrestrial biosphere is thought to have changed since pre-industrial times in response to increasing concentrations of atmospheric CO₂ and variations (anomalies) in inter-annual air temperatures. However, the magnitude of this response, particularly that of various ecosystem types (biomes), is uncertain. Terrestrial carbon models can be used to estimate the direction and size of the terrestrial responses expected, providing that these models have a reasonable theoretical base. We formulated a general model of ecosystem carbon fluxes by linking a process-based canopy photosynthesis model to the Rothamsted soil carbon model for biomes that are not significantly affected by water limitation. The difference between net primary production (NPP) and heterotrophic soil respiration (R_h) represents net ecosystem production (NEP). The model includes (i) multiple compartments for carbon storage in vegetation and soil organic matter, (ii) the effects of seasonal changes in environmental parameters on annual NEP, and (iii) the effects of inter-annual temperature variations on annual NEP. Past, present and projected changes in atmospheric CO₂ concentration and surface air temperature (at different latitudes) were analysed for their effects on annual NEP in tundra, boreal forest and humid tropical forest biomes. In all three biomes, annual NEP was predicted to increase with CO₂ concentration but to decrease with warming. As CO₂ concentrations and temperatures rise, the positive carbon gains through increased NPP are often outweighed by losses through increased R_h, particularly at high latitudes where global warming has been (and is expected to be) most severe. We calculated that, several times during the past 140 years, both the tundra and boreal forest biomes have switched between being carbon sources (annual NEP negative) and being carbon sinks (annual NEP positive). Most recently, significant warming at high latitudes during 1988 and 1990 caused the tundra and boreal forests to be net carbon sources. Humid tropical forests generally have been a carbon sink since 1960. These modelled responses of the various biomes are in agreement with other estimates from either field measurements or geochemical models. Under projected CO₂ and temperature increases, the tundra and boreal forests will emit increasingly more carbon to the atmosphere while the humid tropical forest will continue to store carbon. Our analyses also indicate that the relative increase in the seasonal amplitude of the accumulated NEP within a year is about 0–14% year^?1 for boreal forests and 0–23% year^?1 in the tundra between 1960 and 1990.     

A biome classification of China based on plant functional types and the BIOME3 model

A biome classification for China was established based on plant functional types (PFTs) using the BIOME3 model to include 16 biomes. In the eastern part of China, the PFTs of trees determine mostly the physiognomy of landscape. Biomes range from boreal deciduous coniferous forest/woodland, boreal mixed forest/woodland, temperate mixed forest, temperate broad-leaved deciduous forest, warm-temperate broad-leaved evergreen/mixed forest, warm-temperate/cool-temperate evergreen coniferous forest, xeric woodland/scrub, to tropical seasonal and rain forest, and tropical deciduous forest from north to south. In the northern and western part of China, grass is the dominant PFT. From northeast to west and southwest the biomes range from moist savannas, tall grassland, short grassland, dry savannas, arid shrubland/steppe, desert, to alpine tundra/ice/polar desert. Comparisons between the classification introduced here and the four classifications which were established over the past two decades, i.e. the vegetation classification, the vegetation division, the physical ecoregion, and the initial biome classification have showed that the different aims of biome classifications have resulted in different biome schemes each with its own unique characteristics and disadvantages for global change study. The new biome classification relies not only on climatic variables, but also on soil factor, vegetation functional variables, ecophysiological parameters and competition among the PFTs. It is a comprehensive classification that using multivariables better expresses the vegetation distribution and can be compared with world biome classifications. It can be easily used in the response study of Chinese biomes to global change, regionally and globally.     

Unexpectedly High Bacterial Diversity in Arctic Tundra Relative to Boreal Forest Soils, Revealed by Serial Analysis of Ribosomal Sequence Tags

Arctic tundra and boreal forest soils have globally relevant functions that affect atmospheric chemistry and climate, yet the bacterial composition and diversity of these soils have received little study. Serial analysis of ribosomal sequence tags (SARST) and denaturing gradient gel electrophoresis (DGGE) were used to compare composite soil samples taken from boreal and arctic biomes. This study comprises an extensive comparison of geographically distant soil bacterial communities, involving the analysis of 12,850 ribosomal sequence tags from six composite soil samples. Bacterial diversity estimates were greater for undisturbed arctic tundra soil samples than for boreal forest soil samples, with the highest diversity associated with a sample from an extreme northern location (82^oN). The lowest diversity estimate was obtained from an arctic soil sample that was disturbed by compaction and sampled from a greater depth. Since samples from the two biomes did not form distinct clusters on the basis of SARST data and DGGE fingerprints, factors other than latitude likely influenced the phylogenetic compositions of these communities. The high number of ribosomal sequences analyzed enabled the identification of possible cosmopolitan and endemic bacterial distributions in particular soils.     

Site-level evaluation of satellite-based global terrestrial gross primary production and net primary production monitoring

Operational monitoring of global terrestrial gross primary production (GPP) and net primary production (NPP) is now underway using imagery from the satellite‐borne Moderate Resolution Imaging Spectroradiometer (MODIS) sensor. Evaluation of MODIS GPP and NPP products will require site‐level studies across a range of biomes, with close attention to numerous scaling issues that must be addressed to link ground measurements to the satellite‐based carbon flux estimates. Here, we report results of a study aimed at evaluating MODIS NPP/GPP products at six sites varying widely in climate, land use, and vegetation physiognomy. Comparisons were made for twenty‐five 1 km² cells at each site, with 8‐day averages for GPP and an annual value for NPP. The validation data layers were made with a combination of ground measurements, relatively high resolution satellite data (Landsat Enhanced Thematic Mapper Plus at ～30 m resolution), and process‐based modeling. There was strong seasonality in the MODIS GPP at all sites, and mean NPP ranged from 80 g C m^?2 yr^?1 at an arctic tundra site to 550 g C m^?2 yr^?1 at a temperate deciduous forest site. There was not a consistent over‐ or underprediction of NPP across sites relative to the validation estimates. The closest agreements in NPP and GPP were at the temperate deciduous forest, arctic tundra, and boreal forest sites. There was moderate underestimation in the MODIS products at the agricultural field site, and strong overestimation at the desert grassland and at the dry coniferous forest sites. Analyses of specific inputs to the MODIS NPP/GPP algorithm – notably the fraction of photosynthetically active radiation absorbed by the vegetation canopy, the maximum light use efficiency (LUE), and the climate data – revealed the causes of the over‐ and underestimates. Suggestions for algorithm improvement include selectively altering values for maximum LUE (based on observations at eddy covariance flux towers) and parameters regulating autotrophic respiration.     

Simulation of tree species composition and organic matter accumulation in Finnish boreal forests under changing climatic conditions

A model simulating the regeneration, growth and death of trees and the consequent carbon and nitrogen dynamics of the forest ecosystem was applied to determine the effect of expected temperature rise on tree species composition and the accumulation of organic matter in the boreal forest ecosystem in Finland (between latitudes 60°–70° N). In the southern and middle boreal zones a temperature rise of 2–3° C (temperature for 2 x CO₂) over a period of one hundred years increased the competitive capacity of Scots pine (Pinus sylvestris) and birch species (Betula pendula and B. pubescens), and slowed down the invasion by Norway spruce (Picea abies). In the northern boreal zone a corresponding rise in temperature promoted the invasion of sites by Norway spruce. The accumulation of organic matter was promoted only slightly compared to that taking place in the current climatic conditions.A further doubling of temperature (temperature for 4 x CO₂) over an additional period of two hundred years led to the replacement of coniferous stands with deciduous onesin the southern and middle boreal zones. In the northern boreal zone an admixture of coniferous and deciduous species replaced pure coniferous stands with the latter taking over sites formerly classified as tundra woodland. In the southern and middle boreal zones the replacement of coniferous species induced a substantial decrease in the amount of organic matter; this returned to its former level following the establishment of deciduous species. In the northern boreal zone there was no major change in the amount of organic matter such as occurred in the case of the tundra woodland where the amount of organic matter accumulated was nearly as high as in the northern boreal zone.     

Mid-Holocene and glacial-maximum vegetation geography of the northern continents and Africa

BIOME 6000 is an international project to map vegetation globally at mid‐Holocene (6000 ¹⁴C yr bp ) and last glacial maximum (LGM, 18,000 ¹⁴C yr bp ), with a view to evaluating coupled climate‐biosphere model results. Primary palaeoecological data are assigned to biomes using an explicit algorithm based on plant functional types. This paper introduces the second Special Feature on BIOME 6000. Site‐based global biome maps are shown with data from North America, Eurasia (except South and Southeast Asia) and Africa at both time periods. A map based on surface samples shows the method’s skill in reconstructing present‐day biomes. Cold and dry conditions at LGM favoured extensive tundra and steppe. These biomes intergraded in northern Eurasia. Northern hemisphere forest biomes were displaced southward. Boreal evergreen forests (taiga) and temperate deciduous forests were fragmented, while European and East Asian steppes were greatly extended. Tropical moist forests (i.e. tropical rain forest and tropical seasonal forest) in Africa were reduced. In south‐western North America, desert and steppe were replaced by open conifer woodland, opposite to the general arid trend but consistent with modelled southward displacement of the jet stream. The Arctic forest limit was shifted slighly north at 6000 ¹⁴C yr bp in some sectors, but not in all. Northern temperate forest zones were generally shifted greater distances north. Warmer winters as well as summers in several regions are required to explain these shifts. Temperate deciduous forests in Europe were greatly extended, into the Mediterranean region as well as to the north. Steppe encroached on forest biomes in interior North America, but not in central Asia. Enhanced monsoons extended forest biomes in China inland and Sahelian vegetation into the Sahara while the African tropical rain forest was also reduced, consistent with a modelled northward shift of the ITCZ and a more seasonal climate in the equatorial zone. Palaeobiome maps show the outcome of separate, independent migrations of plant taxa in response to climate change. The average composition of biomes at LGM was often markedly different from today. Refugia for the temperate deciduous and tropical rain forest biomes may have existed offshore at LGM, but their characteristic taxa also persisted as components of other biomes. Examples include temperate deciduous trees that survived in cool mixed forest in eastern Europe, and tropical evergreen trees that survived in tropical seasonal forest in Africa. The sequence of biome shifts during a glacial‐interglacial cycle may help account for some disjunct distributions of plant taxa. For example, the now‐arid Saharan mountains may have linked Mediterranean and African tropical montane floras during enhanced monsoon regimes. Major changes in physical land‐surface conditions, shown by the palaeobiome data, have implications for the global climate. The data can be used directly to evaluate the output of coupled atmosphere‐biosphere models. The data could also be objectively generalized to yield realistic gridded land‐surface maps, for use in sensitivity experiments with atmospheric models. Recent analyses of vegetation‐climate feedbacks have focused on the hypothesized positive feedback effects of climate‐induced vegetation changes in the Sahara/Sahel region and the Arctic during the mid‐Holocene. However, a far wider spectrum of interactions potentially exists and could be investigated, using these data, both for 6000 ¹⁴C yr bp and for the LGM.     

Modelling the vegetation of China using the process-based equilibrium terrestrial biosphere model BIOME3

1 We model the potential vegetation and annual net primary production (NPP) of China on a 10′ grid under the present climate using the processed‐based equilibrium terrestrial biosphere model BIOME3. The simulated distribution of the vegetation was in general in good agreement with the potential natural vegetation based on a numerical comparison between the two maps using the ΔV statistic (ΔV = 0.23). Predicted and measured NPP were also similar, especially in terms of biome‐averages. 2 A coupled ocean–atmosphere general circulation model including sulphate aerosols was used to drive a double greenhouse gas scenario for 2070–2099. Simulated vegetation maps from two different CO₂ scenarios (340 and 500 p.p.m.v.) were compared to the baseline biome map using ΔV. Climate change alone produced a large reduction in desert, alpine tundra and ice/polar desert, and a general pole‐ward shift of the boreal, temperate deciduous, warm–temperate evergreen and tropical forest belts, a decline in boreal deciduous forest and the appearance of tropical deciduous forest. The inclusion of CO₂ physiological effects led to a marked decrease in moist savannas and desert, a general decrease for grasslands and steppe, and disappearance of xeric woodland/scrub. Temperate deciduous broadleaved forest, however, shifted north to occupy nearly half the area of previously temperate mixed forest. 3 The impact of climate change and increasing CO₂ is not only on biogeography, but also on potential NPP. The NPP values for most of the biomes in the scenarios with CO₂ set at 340 p.p.m.v. and 500 p.p.m.v. are greater than those under the current climate, except for the temperate deciduous forest, temperate evergreen broadleaved forest, tropical rain forest, tropical seasonal forest, and xeric woodland/scrub biomes. Total vegetation and total carbon is simulated to increase significantly in the future climate scenario, both with and without the CO₂ direct physiological effect. 4 Our results show that the global process‐based equilibrium terrestrial biosphere model BIOME3 can be used successfully at a regional scale.     

7. Tundra stream macroalgae of North America: composition, distribution and physiological adaptations

Eighty-three infrageneric taxa of stream macroalgae have been reported from tundra regions of North America, composed of 32 cyanobacteria, 35 Chlorophyta, 10 Chrysophyta and 6 Rhodophyta. There are few if any endemics represented in this flora. The most widespread species are the cyanobacteria Rivularia minutula, Nostoc commune and Tolypothrix tenuis as well as asexual populations of the chlorophyte genus Zygnema. The relative contribution of cyanobacteria compared to that of the Chlorophyta increases from the low to high arctic. Number of species per segment ranges from 0 to 7, with a mean of 2.8, and varies little between the low and high arctic. The percentage of stream bottom covered by macroalgae ranges from 0 to ca. 75%; mean cover values for low and high arctic streams are ca. 12 and 8%, respectively. Tundra macroalgae tend to be more abundant and diverse in less rigorously flowing stream sections. Most species tolerate prolonged freezing by forming resistant vegetative cells with thick walls, plentiful reserves and low molecular weight solutes to lower the freezing point. Many tundra stream macroalgae also produce ‘sunscreen’ pigments to reduce exposure to damaging radiation in the blue and ultraviolet regions. Nutrients tend to be low and phosphorus is often limiting in these systems. Arctic streams appear to differ from those of Antarctica in having potential grazers of macroalgae, such as the chironomid Diamesa, the mayfly Baetis and the caddisfly Brachycentrus.     

Unexpectedly high bacterial diversity in arctic tundra relative to boreal forest soils, revealed by serial analysis of ribosomal sequence tags

Arctic tundra and boreal forest soils have globally relevant functions that affect atmospheric chemistry and climate, yet the bacterial composition and diversity of these soils have received little study. Serial analysis of ribosomal sequence tags (SARST) and denaturing gradient gel electrophoresis (DGGE) were used to compare composite soil samples taken from boreal and arctic biomes. This study comprises an extensive comparison of geographically distant soil bacterial communities, involving the analysis of 12,850 ribosomal sequence tags from six composite soil samples. Bacterial diversity estimates were greater for undisturbed arctic tundra soil samples than for boreal forest soil samples, with the highest diversity associated with a sample from an extreme northern location (82(o)N). The lowest diversity estimate was obtained from an arctic soil sample that was disturbed by compaction and sampled from a greater depth. Since samples from the two biomes did not form distinct clusters on the basis of SARST data and DGGE fingerprints, factors other than latitude likely influenced the phylogenetic compositions of these communities. The high number of ribosomal sequences analyzed enabled the identification of possible cosmopolitan and endemic bacterial distributions in particular soils.     

Helminth parasites of wolves (Canis lupus): a species list and an analysis of published prevalence studies in Nearctic and Palaearctic populations

A literature survey was undertaken in order to draw up a definitive list of helminth parasites of the wolf, Canis lupus. From 27 papers a total of 72 helminth species from 40 genera were recorded that infect wolves, of which 93% were identified from the gastrointestinal tract at necropsy. They comprised 28 species of nematode, 27 species of cestode, 16 species of trematode and one acanthocephalan. Of these, 46 species were able to be included in further meta-analysis of prevalence data derived from 25 publications for which the total number of wolves examined was 1282 (1066 from Nearctic populations, and 216 from the Palaearctic region). These two populations were further subdivided into three relevent ecosystems or biomes, i.e. temperate/montane (n=216), boreal (n=805) or tundra (n=261). The meta-analysis of relative prevalence indicated the most common helminth species to be the tapeworm Taenia hydatigena, which occurred at relative rates of >30% for either zoogeographic region as well as in each of the three biomes. The related tapeworm, Echinococcus granulosus also exhibited high meta-prevalence (>19%) in all host biomes. The hookworm Uncinaria stenocephala was the most prevalent nematode species by meta-analysis (meta-prevalence 44.9%) in the temperate/montane biome, while the ascarid Toxascaris leonina was the dominant helminth species (meta-prevalence 73.9%) in the tundra wolf populations. Trematodes in the genus Alaria were the dominant fluke (meta-prevalence 3-5%) in all biomes. Analysis of published studies for helminth biodiversity using the Shannon-Wiener index based on species number and meta-prevalence by region or biome, indicated that highest helminth diversity occurred in wolf populations of the temperate/ montane biome (Palaearctic), and was lowest in tundra wolf populations of the Nearctic (P<0.05). Helminth species assemblage in European wolf populations was therefore at least as great or more varied than was recorded for the larger less disturbed wolf populations of North America.     

A meta‐analysis of declines in local species richness from human disturbances

There is high uncertainty surrounding the magnitude of current and future biodiversity loss that is occurring due to human disturbances. Here, we present a global meta‐analysis of experimental and observational studies that report 327 measures of change in species richness between disturbed and undisturbed habitats across both terrestrial and aquatic biomes. On average, human‐mediated disturbances lead to an 18.3% decline in species richness. Declines in species richness were highest for endotherms (33.2%), followed by producers (25.1%), and ectotherms (10.5%). Land‐use change and species invasions had the largest impact on species richness resulting in a 24.8% and 23.7% decline, respectively, followed by habitat loss (14%), nutrient addition (8.2%), and increases in temperature (3.6%). Across all disturbances, declines in species richness were greater for terrestrial biomes (22.4%) than aquatic biomes (5.9%). In the tropics, habitat loss and land‐use change had the largest impact on species richness, whereas in the boreal forest and Northern temperate forests, species invasions had the largest impact on species richness. Along with revealing trends in changes in species richness for different disturbances, biomes, and taxa, our results also identify critical knowledge gaps for predicting the effects of human disturbance on Earth's biomes.     

Distribution of Selected Plant Parasitic Nematodes Relative to Vegetation and Edaphic Factors

The occurrence of selected plant-parasitic nematodes in the hemlock-hardwood-white pine, boreal forest, tundra, and oak-hickory associations in some northern states was compared. Helicotylenchus platyurus and Xiphinema americanum were not found in the boreal forest and tundra, and occurred infrequently in the hemlock-hardwood-white pine areas. They were found frequently, however, in the oak-hickory forest of Iowa. It is questioned that vegetational differences among the areas account directly for the major differences in nematode occurrence. Presence and absence of nematodes and their numbers in the oak-hickory association were clustered by similarity coefficients by sites and correlated with soil pH, percentage organic matter, percentage sand-silt-clay, and field capacity. Of the soil factors measured, pH gave the strongest correlations with nematode numbers. Xiphinema chambersi was found only in soils with a pH between 4.5 and 6.4 while the largest numbers of H. platyurus, H. pseudorobustus, and X. americanum occurred in soil above pH 6.0.     

森林土壤有机碳深度分布模型的构建与应用

了解森林土壤有机碳(SOC)的深度分布模式对正确估算森林碳储量,充分发挥森林碳汇功能,减缓全球气候变化有着重要意义。选取寒温带针叶林、温带落叶林、亚热带针阔混交林、热带常绿阔叶林等4类森林生物群系,建立SOC深度分布数据库,构建SOC质量密度的深度分布模型;使用Nash-Sutcliffe效率系数(E)、误差百分比(PE)、决定系数(R~2)等统计参量评定模型的模拟效果;利用构建的深度分布模型外推更深层SOC密度。研究结果表明:(1)本文所构建的森林SOC深度分布模型模拟值与观测值较为吻合,Nash-Sutcliffe效率E、误差百分比PE和决定系数R~2平均为0.74、6.95%、0.88(P0.05),模型模拟能力较高(E0.6),模拟误差值低于可接受的临界值(PE±15%),说明构建的模型可以对该地区森林SOC密度值进行估算;(2)寒温带针叶林0—20 cm层SOC质量密度较高,热带常绿阔叶林较低;20 cm以下则是寒温带针叶林较低,热带常绿阔叶林较高,热带常绿阔叶林具有更深层的SOC分布;用0—100 cm深度的SOC来表征区域SOC储量时结果偏低。若考虑0—200 cm深度,0—100 cm深度SOC值平均偏低约21.8%,在热带地区这种偏低趋势可能更加突出,误差可能更大。(3)模型对表层SOC密度有偏低预测趋势,对深层SOC密度预测值可能偏高;作为一个森林SOC深度分布模拟工具,模型可以在有限区域条件下估算不同深度SOC密度值。     

Checkerboard score–area relationships reveal spatial scales of plant community structure

Identifying the spatial scale at which particular mechanisms influence plant community assembly is crucial to understanding the mechanisms structuring communities. It has long been recognized that many elements of community structure are sensitive to area; however the majority of studies examining patterns of community structure use a single relatively small sampling area. As different assembly mechanisms likely cause patterns at different scales we investigate how plant species co‐occurrence patterns change with sampling unit scale. We use the checkerboard score as an index of species segregation, and examine species C‐score1–sampling area patterns in two ways. First, we show via numerical simulation that the C‐score–area relationship is necessarily hump shaped with respect to sample plot area. Second we examine empirical C‐score–area relationships in arctic tundra, grassland, boreal forest and tropical forest communities. The minimum sampling scale where species co‐occurrence patterns were significantly different from the null model expectation was at 0.1 m² in the tundra, 0.2 m² in grassland, and 0.2 ha in both the boreal and tropical forests. Species were most segregated in their co‐occurrence (maximum C‐score) at 0.3 m² in the tundra (0.54 3 0.54 m quadrats), 1.5 m² in the grassland (1.2 3 1.2 m quadrats), 0.26 ha in the tropical forest (71 3 71 m quadrats), and a maximum was not reached at the largest sampling scale of 1.4 ha in the boreal forest. The most important finding is that the dominant scales of community structure in these systems are large relative to plant body size, and hence we infer that the dominant mechanisms structuring these communities must be at similarly large scales. This provides a method for identifying the spatial scales at which communities are maximally structured; ecologists can use this information to develop hypotheses and experiments to test scale‐specific mechanisms that structure communities.     

Global pattern and controls of biological nitrogen fixation under nutrient enrichment: A meta‐analysis

Biological nitrogen (N) fixation (BNF), an important source of N in terrestrial ecosystems, plays a critical role in terrestrial nutrient cycling and net primary productivity. Currently, large uncertainty exists regarding how nutrient availability regulates terrestrial BNF and the drivers responsible for this process. We conducted a global meta‐analysis of terrestrial BNF in response to N, phosphorus (P), and micronutrient (Micro) addition across different biomes (i.e, tropical/subtropical forest, savanna, temperate forest, grassland, boreal forest, and tundra) and explored whether the BNF responses were affected by fertilization regimes (nutrient‐addition rates, duration, and total load) and environmental factors (mean annual temperature [MAT], mean annual precipitation [MAP], and N deposition). The results showed that N addition inhibited terrestrial BNF (by 19.0% (95% confidence interval [CI]: 17.7%?20.3%); hereafter), Micro addition stimulated terrestrial BNF (30.4% [25.7%?35.3%]), and P addition had an inconsistent effect on terrestrial BNF, i.e., inhibiting free‐living N fixation (7.5% [4.4%?10.6%]) and stimulating symbiotic N fixation (85.5% [25.8%?158.7%]). Furthermore, the response ratios (i.e., effect sizes) of BNF to nutrient addition were smaller in low‐latitude (<30°) biomes (8.5%?36.9%) than in mid‐/high‐latitude (≥30°) biomes (32.9%?61.3%), and the sensitivity (defined as the absolute value of response ratios) of BNF to nutrients in mid‐/high‐latitude biomes decreased with decreasing latitude (p ≤ 0.009; linear/logarithmic regression models). Fertilization regimes did not affect this phenomenon (p > 0.05), but environmental factors did affect it (p < 0.001) because MAT, MAP, and N deposition accounted for 5%?14%, 10%?32%, and 7%?18% of the variance in the BNF response ratios in cold (MAT < 15°C), low‐rainfall (MAP < 2,500 mm), and low‐N‐deposition (<7 kg ha^?1 year^?1) biomes, respectively. Overall, our meta‐analysis depicts a global pattern of nutrient impacts on terrestrial BNF and indicates that certain types of global change (i.e., warming, elevated precipitation and N deposition) may reduce the sensitivity of BNF in response to nutrient enrichment in mid‐/high‐latitude biomes.     

Regional‐scale directional changes in abundance of tree species along a temperature gradient in Japan

Climate changes are assumed to shift the ranges of tree species and forest biomes. Such range shifts result from changes in abundances of tree species or functional types. Owing to global warming, the abundance of a tree species or functional type is expected to increase near the colder edge of its range and decrease near the warmer edge. This study examined directional changes in abundance and demographic parameters of forest trees along a temperature gradient, as well as a successional gradient, in Japan. Changes in the relative abundance of each of four functional types (evergreen broad‐leaved, deciduous broad‐leaved, evergreen temperate conifer, and evergreen boreal conifer) and the demography of each species (recruitment rate, mortality, and population growth rate) were analyzed in 39 permanent forest plots across the Japanese archipelago. Directional changes in the relative abundance of functional types were detected along the temperature gradient. Relative abundance of evergreen broad‐leaved trees increased near their colder range boundaries, especially in secondary forests, coinciding with the decrease in deciduous broad‐leaved trees. Similarly, relative abundance of deciduous broad‐leaved trees increased near their colder range boundaries, coinciding with the decrease in boreal conifers. These functional‐type‐level changes were mainly due to higher recruitment rates and partly to the lower mortality of individual species at colder sites. This is the first report to show that tree species abundances in temperate forests are changing directionally along a temperature gradient, which might be due to current or past climate changes as well as recovery from past disturbances.     

The greening and browning of Alaska based on 1982–2003 satellite data

Aim To examine the trends of 1982–2003 satellite‐derived normalized difference vegetation index (NDVI) values at several spatial scales within tundra and boreal forest areas of Alaska. Location Arctic and subarctic Alaska. Methods Annual maximum NDVI data from the twice monthly Global Inventory Modelling and Mapping Studies (GIMMS) NDVI 1982–2003 data set with 64‐km² pixels were extracted from a spatial hierarchy including three large regions: ecoregion polygons within regions, ecozone polygons within boreal ecoregions and 100‐km climate station buffers. The 1982–2003 trends of mean annual maximum NDVI values within each area, and within individual pixels, were computed using simple linear regression. The relationship between NDVI and temperature and precipitation was investigated within climate station buffers. Results At the largest spatial scale of polar, boreal and maritime regions, the strongest trend was a negative trend in NDVI within the boreal region. At a finer scale of ecoregion polygons, there was a strong positive NDVI trend in cold arctic tundra areas, and a strong negative trend in interior boreal forest areas. Within boreal ecozone polygons, the weakest negative trends were from areas with a maritime climate or colder mountainous ecozones, while the strongest negative trends were from warmer basin ecozones. The trends from climate station buffers were similar to ecoregion trends, with no significant trends from Bering tundra buffers, significant increasing trends among arctic tundra buffers and significant decreasing trends among interior boreal forest buffers. The interannual variability of NDVI among the arctic tundra buffers was related to the previous summer warmth index. The spatial pattern of increasing tundra NDVI at the pixel level was related to the west‐to‐east spatial pattern in changing climate across arctic Alaska. There was no significant relationship between interannual NDVI and precipitation or temperature among the boreal forest buffers. The decreasing NDVI trend in interior boreal forests may be due to several factors including increased insect/disease infestations, reduced photosynthesis and a change in root/leaf carbon allocation in response to warmer and drier growing season climate. Main conclusions There was a contrast in trends of 1982–2003 annual maximum NDVI, with cold arctic tundra significantly increasing in NDVI and relatively warm and dry interior boreal forest areas consistently decreasing in NDVI. The annual maximum NDVI from arctic tundra areas was strongly related to a summer warmth index, while there were no significant relationships in boreal areas between annual maximum NDVI and precipitation or temperature. Annual maximum NDVI was not related to spring NDVI in either arctic tundra or boreal buffers.     

Changes in the structure and function of northern Alaskan ecosystems when considering variable leaf‐out times across groupings of species in a dynamic vegetation model

The phenology of arctic ecosystems is driven primarily by abiotic forces, with temperature acting as the main determinant of growing season onset and leaf budburst in the spring. However, while the plant species in arctic ecosystems require differing amounts of accumulated heat for leaf‐out, dynamic vegetation models simulated over regional to global scales typically assume some average leaf‐out for all of the species within an ecosystem. Here, we make use of air temperature records and observations of spring leaf phenology collected across dominant groupings of species (dwarf birch shrubs, willow shrubs, other deciduous shrubs, grasses, sedges, and forbs) in arctic and boreal ecosystems in Alaska. We then parameterize a dynamic vegetation model based on these data for four types of tundra ecosystems (heath tundra, shrub tundra, wet sedge tundra, and tussock tundra), as well as ecotonal boreal white spruce forest, and perform model simulations for the years 1970–2100. Over the course of the model simulations, we found changes in ecosystem composition under this new phenology algorithm compared with simulations with the previous phenology algorithm. These changes were the result of the differential timing of leaf‐out, as well as the ability for the groupings of species to compete for nitrogen and light availability. Regionally, there were differences in the trends of the carbon pools and fluxes between the new phenology algorithm and the previous phenology algorithm, although these differences depended on the future climate scenario. These findings indicate the importance of leaf phenology data collection by species and across the various ecosystem types within the highly heterogeneous Arctic landscape, and that dynamic vegetation models should consider variation in leaf‐out by groupings of species within these ecosystems to make more accurate projections of future plant distributions and carbon cycling in Arctic regions.