Stable isotopic composition of carbon and nitrogen and nitrogen content in vascular epiphytes along an altitudinal transect*

The foliar content of nitrogen and the relative abundances of ¹³C and ¹⁵N were analysed in vascular epiphytes collected from six sites along an altitudinal gradient from tropical dry forests to humid montane forests in eastern Mexico. The proportion of epiphyte species showing crassulacean acid metabolism (CAM) (atmospheric bromeliads, thick-leaved orchids, Cactaceae, and Crassulaceae) decreased with increasing elevation and precipitation from 58 to 6%. Atmospheric bromeliads, almost all of which had δ ¹³C values indicating CAM, were more depleted in ¹⁵N (x = ? 10·9‰ ± 2·11) than the C₃ bromeliads which form water-storing tanks ( ? 6·05‰ ± 2·26). As there was no difference in δ ¹⁵N values between C₃ and CAM orchids, the difference in bromeliads was not related to photosynthetic pathways but to different nitrogen sources. While epiphytes with strong ¹⁵N depletion appear to obtain their nitrogen mainly from direct atmospheric deposition, others have access to nitrogen in intercepted water and from organic matter decomposing on branches and in their phytotelmata. Bromeliads and succulent orchids had a lower foliar nitrogen content than thin-leaved orchids, ferns and Piperaceae. Ground-rooted hemi-epiphytes exhibited the highest nitrogen contents and δ ¹⁵N values.     

Tissue-specific variation of δC in mature canopy trees in a temperate forest in central Europe

Stable carbon isotope analysis has become a key tool in functional ecology, yet considerable natural variability often limits the interpretations. In this study we document the spatial, taxonomic, temporal and tissue-specific δ¹³C variability in 10 tree species of a temperate European forest. The Swiss Canopy Crane provided access to the three dimensional space within 55 trees 30–35 m high representing the genera Acer, Carpinus, Fagus, Prunus, Quercus, Tilia, Abies, Larix, Picea and Pinus. The results from six broad-leaved and four conifer species (seven deciduous, three evergreen) documented that the species effect was not significant in contrast to tissue-specific and spatial differences in the canopy. Year-to-year differences were not large but still significant.

Our analysis confirmed a significant difference between δ¹³C of foliage collected in the upper and lower canopy, but revealed no systematic differences with respect to azimuthal directions in tree crowns of the broad-leaved trees, as opposed to the conifers, which show clear differences between the sun-exposed and the shaded side. Tissue-specific differences were significant, despite surprisingly similar mean values for most tissue types. Such tissue effect was largely due to young branch xylem, which exhibited a systematic less negative deviation from the other tissue types. These findings were consistent across the species tested and provided some guidelines towards a representative sampling strategy for ¹³C analysis.     

Root dynamics influence tree–grass coexistence in an Australian savanna

Both resource and disturbance controls have been invoked to explain tree persistence among grasses in savannas. Here we determine the extent to which competition for available resources restricts the rooting depth of both grasses and trees, and how this may influence nutrient cycling under an infrequently burned savanna near Darwin, Australia. We sampled fine roots <2 mm in diameter from 24 soil pits under perennial as well as annual grasses and three levels of canopy cover. The relative proportion of C₃ (trees) and C₄ (grasses) derived carbon in a sample was determined using mass balance calculations. Our results show that regardless of the type of grass both tree and grass roots are concentrated in the top 20 cm of the soil. While trees have greater root production and contribute more fine root biomass grass roots contribute a disproportional amount of nitrogen and carbon to the soil relative to total root biomass. We postulate that grasses maintain soil nutrient pools and provide biomass for regular fires that prevent forest trees from establishing while savanna trees, are important for increasing soil N content, cycling and mineralization rates. We put forward our ideas as a hypothesis of resource‐regulated tree–grass coexistence in tropical savannas.     

Present dynamics of the savanna-forest boundary in the Congolese Mayombe: a pedological,botanical and isotopic (13C and 14C) study

Isolated savannas enclosed by forest are especially abundant in the eastern part of the Congolese Mayombe. They are about 3000 years old, and were more extensive some centuries ago. The boundary between forest and savanna is very abrupt, as a consequence of the numerous savanna fires lit by hunters. Floristic composition and vegetation structure data, organic carbon ratios, ¹⁴C and ¹³C measurements presented here show that forest is spreading over savanna at the present time and suggest that the rate of forest encroachment is is currently between 14 and 75 m per century, and more probably about 20–50 m per century. As most savannas are less than 1 km across, such rates mean, assuming there are no changes in environmental conditions, that enclosed savannas could completely disappear in the Mayombe in about 1000–2000 years.     

Broad‐scale vegetation‐environment relationships in Eurasian high‐latitude areas

Question: How is tundra vegetation related to climatic, soil chemical, geological variables and grazing across a very large section of the Eurasian arctic area? We were particularly interested in broad‐scale vegetation‐environment relationships and how well do the patterns conform to climate‐vegetation schemes. Material and Methods: We sampled vegetation in 1132 plots from 16 sites from different parts of the Eurasian tundra. Clustering and ordination techniques were used for analysing compositional patterns. Vegetation‐environment relationships were analysed by fitting of environmental vectors and smooth surfaces onto non‐metric multidimensional scaling scattergrams. Results: Dominant vegetation differentiation was associated with a complex set of environmental variables. A general trend differentiated cold and continental areas from relatively warm and weakly continental areas, and several soil chemical and physical variables were associated with this broad‐scaled differentiation. Especially soil chemical variables related to soil acidity (pH, Ca) showed linear relationships with the dominant vegetation gradient. This was closely related to increasing cryoperturbation, decreasing precipitation and cooler conditions. Remarkable differences among relatively adjacent sites suggest that local factors such as geological properties and lemming grazing may strongly drive vegetation differentiation. Conclusions: Vegetation differentiation in tundra areas conforms to a major ecocline underlain by a complex set of environmental gradients, where precipitation, thermal conditions and soil chemical and physical processes are coupled. However, local factors such as bedrock conditions and lemming grazing may cause marked deviations from the general climate‐vegetation models. Overall, soil chemical factors (pH, Ca) turned out to have linear relationship with the broad‐scale differentiation of arctic vegetation.     

Digitized photographs in vegetation analysis ‐ a comparison of cover estimates

Abstract. The use of digital images in measuring plant species cover and species number in boreal forest vegetation was studied. Plant cover was estimated by manual delineation on photographs and subsequent digitized measurement of the areas. This value was regarded as the reference and compared with cover obtained by visual estimate, point frequency and automatic image analysis methods. The automatic image analysis was based on scanned photographs. Supervised image classification with ERDAS software was then used to distinguish between the covers of different plant species. When comparing the ability of the methods to detect species, the visual estimate method gave values similar to the reference. The study material was collected from three sites along a heavy‐metal pollution transect in western Finland. All four methods detected, in a similar manner, differences between the plant species abundances along the transect. Compared with the reference, the digital images underestimated the cover of the lichens Cladina spp. and Cetraria islandica, but gave similar estimates for the dwarf shrubs Empetrum nigrum and Vaccinium vitis‐idaea. The point frequency method overestimated the cover of all the species studied. The visual estimates of lichens were close to the reference, while the dwarf shrub covers were overestimated. The number of species detected using supervised image analysis and the point frequency method was lower than that with visual estimation. Visual estimation was faster, and the estimate closer to the reference cover values than the others. Digital images may be useful in detecting changes in some selected species in vegetation with a simple vertical structure but with taller, multilayered vegetation and a higher species number, the reliability of the cover estimates is lower.     

Seed bank dynamics in tall‐tussock grasslands along an altitudinal gradient

Abstract. We studied the germinable soil seed bank of tall‐tussock grasslands along an altitudinal gradient in the mountains of central Argentina. We selected 10 sampling plots at three altitudinal levels (1200 m, 1600 m and 2200 m). We assessed the composition of the established vegetation and took ten compound soil samples (0 ‐ 5 cm depth) at each plot in autumn and spring. The soil samples were sieved, chilled, and incubated in a glasshouse to assess the composition of the seed bank. The similarity between the composition of the seed bank flora and that of the established vegetation was low throughout the gradient. Most species did not change their seed bank strategy along the gradient. Seed bank richness and density increased with altitude. Most species had a persistent seed bank at all altitudinal levels, and the proportion of such species increased with altitude. These results suggest that a cold climate directly and/or indirectly favours the formation of seed banks and seed persistence in the soil.     

Soil as a mediator in plant‐plant interactions in a semi‐arid community

Abstract. Competition and facilitation may occur simultaneously in plant communities, and the prevalence of either process depends on abiotic conditions. Here we attempt a community‐wide approach in the analysis of plant interactions, exploring whether in a semi‐arid environment positive or negative interactions predominate and whether there are differences among co‐occurring shrub species. Most shrubs in our plot exerted significant effects on their understorey communities, ranging from negative to positive. We found a clear case of interference and another case where the effect was neutral, but facilitation predominated and the biomass of annuals under most shrubs in our community was larger than in gaps. Effects on soil water and fertility were revealed as the primary source of facilitation; the build‐up of soil organic matter changed soil physical properties and improved soil water relations. Facilitation by shrubs involved decoupling of soil temperature and moisture. Sheltering from direct radiation had an effect on productivity, but significant differences in understorey biomass did not parallel understorey light environment. A positive balance of the interaction among plants, essentially mediated by changes in soil properties, is the predominant outcome of plant interactions in this semi‐arid community.     

长期垄作稻田腐殖质稳定碳同位素丰度(δ13C)分布特征

研究了四川盆地丘陵区连续16年垄（宽垄）作稻田土壤稳定碳库腐殖质组分的稳定碳同位素(δ¹³C)分布特征.结果表明： 稻田土壤有机碳含量为宽垄作＞垄作＞水旱轮作.腐殖质碳以胡敏素为主,占土壤碳含量的21%～30%,提取碳以胡敏酸为主,分别占土壤有机碳和腐殖质的17%～21%和38%～65%.土壤有机碳的δ¹³C值介于-27.9‰～-25.6‰,20～40 cm和0～5 cm土壤有机碳δ¹³C值之差约为1.9‰.土壤胡敏酸δ¹³C值比土壤有机碳低1‰～2‰,更接近于油菜和水稻秸秆及根系的δ¹³C值.土壤富里酸δ¹³C值分别较土壤有机碳和胡敏酸高2‰和4‰.耕作层和犁底层胡敏素δ¹³C值分别介于-23.7‰～-24.9‰和-22.6‰～-24.2‰,δ¹³C值的变化反映了耕层中腐殖质的新老混合现象.各有机组分δ¹³C值递减顺序为：胡敏素＞富里酸＞土壤有机碳＞稻草（油菜）残体＞胡敏酸.长期水稻种植有利于增加土壤有机碳含量,同时,耕作方式影响土壤腐殖质δ¹³C在耕作层和犁底层中的分布格局.     

Colonization dynamics and facilitative impacts of a nitrogen‐fixing shrub in primary succession

Abstract. Whether nitrogen‐fixing plants facilitate or inhibit species change in primary succession is best resolved by examining their impacts throughout the plant's entire life cycle from arrival to senescence. We experimentally examined two aspects of the successional impacts of a nitrogen‐fixing shrub, Coriaria arborea, on Mt. Tarawera, a volcano in New Zealand: factors limiting Coriaria colonization and impacts of Coriaria‐induced soil changes on a later successional tree, Griselinia littoralis. Coriaria germination was promoted by artificial wind protection and by the presence of heath shrubs. Transplanted Coriaria seedlings survived only if nodulated with Frankia, and the addition of Coriaria‐enriched soils slowed Coriaria seedling growth and did not improve seedling survival. This explained why Coriaria seedlings were found mostly in protected habitats away from adult Coriaria, and suggested that Coriaria thickets are not self‐replacing. Coriaria increased soil fertility by developing a 4 cm thick organic soil horizon that was richer in nitrogen (tenfold) and phosphorus (threefold) than pre‐Coriaria stages. These soil changes resulted in three‐ to sixfold increases in growth of Griselinia when it was grown in Coriaria‐enriched soils in a glasshouse. Coriaria's net effect on primary succession is facilitative, but the establishment of Coriaria is itself facilitated by the amelioration of the physical habitat by earlier colonists, suggesting facilitation is important throughout the life cycle of Coriaria. Sequential facilitative events determine the order of species replacements in this study but inhibition, linked to the developmental stages of Coriaria, may determine the rate of species change.     

Long‐term dynamics in a planted conifer forest with spontaneous ingrowth of broad‐leaved trees

Question: What are the age structure and growth trends in a 160‐year old not‐managed Pinus sylvestris plantation with spontaneous development of Quercus robur and can recruitment of Q. robur be related to the radial growth pattern of the P. sylvestris overstorey? Location: Mattemburgh forest reserve, The Netherlands. Methods: Throughout the forest, we sampled 103 oaks and 102 pines with an increment corer. Tree ring widths were measured and cross‐dated to produce mean ring width series. With these data we determined tree ages, investigated growth trends and identified growth releases and suppressions. Results: Q. robur is uneven‐aged: some individuals recruited around 1925, but most reached coring height in the 1940s. The latter recruitment period related to a transition from stressed to released growth of the overstorey pines, growth releases of the oldest Q. robur and occurrence of P. sylvestris regeneration. No further recruitment has taken place since 1950. Conclusions: This study demonstrates that an old pine plantation can develop spontaneously into well‐structured pine forest with an understorey of oak and pine. However, understorey recruitment in these forest types is not a continuous process and in this case a single allogenic canopy disturbance triggered its establishment.     

Trade‐offs in soil carbon protection mechanisms under aerobic and anaerobic conditions

Oxygen (O₂) limitation is generally understood to suppress oil carbon (C) decomposition and is a key mechanism impacting terrestrial C stocks under global change. Yet, O₂ limitation may differentially impact kinetic or thermodynamic versus physicochemical C protection mechanisms, challenging our understanding of how soil C may respond to climate‐mediated changes in O₂ dynamics. Although O₂ limitation may suppress decomposition of new litter C inputs, release of physicochemically protected C due to iron (Fe) reduction could potentially sustain soil C losses. To test this trade‐off, we incubated two disparate upland soils that experience periodic O₂ limitation—a tropical rainforest Oxisol and a temperate cropland Mollisol—with added litter under either aerobic (control) or anaerobic conditions for 1 year. Anoxia suppressed total C loss by 27% in the Oxisol and by 41% in the Mollisol relative to the control, mainly due to the decrease in litter‐C decomposition. However, anoxia sustained or even increased decomposition of native soil‐C (11.0% vs. 12.4% in the control for the Oxisol and 12.5% vs. 5.3% in the control for the Mollisol, in terms of initial soil C mass), and it stimulated losses of metal‐ or mineral‐associated C. Solid‐state ¹³C nuclear magnetic resonance spectroscopy demonstrated that anaerobic conditions decreased protein‐derived C but increased lignin‐ and carbohydrate‐C relative to the control. Our results indicate a trade‐off between physicochemical and kinetic/thermodynamic C protection mechanisms under anaerobic conditions, whereby decreased decomposition of litter C was compensated by more extensive loss of mineral‐associated soil C in both soils. This challenges the common assumption that anoxia inherently protects soil C and illustrates the vulnerability of mineral‐associated C under anaerobic events characteristic of a warmer and wetter future climate.     

Vegetation patterns in heterogeneous landscapes: The importance of history and environment

Abstract. Throughout the eastern United States, plant species distributions and community patterns have developed in response to heterogeneous environmental conditions and a wide range of historical factors, including complex histories of natural and anthropogenic disturbance. Despite increased recognition of the importance of disturbance in determining forest composition and structure, few studies have assessed the relative influence of current environment and historical factors on modern vegetation, in part because detailed knowledge of prior disturbance is often lacking. In the present study, we investigate modern and historical factors that control vegetation patterns at Harvard Forest in central Massachusetts, USA. Similar to the forested uplands throughout the northeastern United States, the site is physiographically heterogeneous and has a long and complex history of natural and anthropogenic disturbance. However, data on forest composition and disturbance history collected over the past > 90 years allow us to evaluate the importance of historical factors rigorously, which is rarely possible on other sites. Soil analyses and historical sources document four categories of historical land use on areas that are all forested today: cultivated fields, improved pastures/mowings, unimproved pastures, and continuously forested woodlots. Ordination and logistic regressions indicate that although species have responded individualistically to a wide range of environmental and disturbance factors, many species are influenced by three factors: soil drainage, land use history, and C:N ratios. Few species vary in accordance with ionic gradients, damage from the 1938 hurricane, or a 1957 fire. Contrary to our expectation that the effects of disturbance will diminish over time, historical land use predicts 1992 vegetation composition better than 1937 composition, perhaps because historical woodlots have become increasingly differentiated from post-agricultural stands through the 20th century. Interpretations of modern vegetation must consider the importance of historical factors in addition to current environmental conditions. However, because disturbances such as land use practices and wind damage are complex, it is often difficult to detect disturbance effects using multivariate approaches, even when the broad history of disturbance is known.     

Formation and maintenance of community boundaries in a sub‐alpine forest landscape in northern Japan

Abstract. We focused on community boundaries in a sub‐alpine forest landscape in the Shiretoko Peninsula, northern Japan. Gradient‐directed transects were conducted on the northwestern slope (ranging 500–600 m a.s.l.) of Mount On'nebetsu (1331 m), where complex topography was formed by past landslides. Pioneer Picea glehnii made up a mosaic of pure stands related to landslides. Structural and compositional changes from P. glehnii pure stands to P. glehnii and Abies sachalinensis mixed stands were characterized by ca. 20 m transitional zones over the landscape. Stand density of the species changed across boundaries. A. sachalinensis preferred less undulated slopes with deep soil and P. glehnii preferred undulated rocky sites. Positive spatial associations between overstorey‐understorey P. glehnii were found at undulated core parts of P. glehnii pure stands. Short‐lived A. sachalinensis grew faster to the smaller maximum size than long‐lived P. glehnii. Undulated topography controlled the increase of A. sachalinensis and provided regeneration sites for P. glehnii, which prevented the general trend of canopy replacement from P. glehnii to late‐successional A. sachalinensis. However, the locations of current boundaries were not accordant with the topographic changes in the meso‐scale landscape. Initial P. glehnii pure stands would extend to larger areas if current patterns reflect vegetation recovery since the last landslide. P. glehnii pure stands with accurate boundaries were not maintained by topographic complexity, but were dynamically arranged by the one‐sided canopy replacements from P. glehnii to A. sachalinensis at less undulated slopes in the sub‐alpine forest landscape.     

Carbon sequestration and turnover in soil under the energy crop Miscanthus: repeated 13C natural abundance approach and literature synthesis

The stability and turnover of soil organic matter (SOM) are a very important but poorly understood part of carbon (C) cycling. Conversion of C₃ grassland to the C₄ energy crop Miscanthus provides an ideal opportunity to quantify medium‐term SOM dynamics without disturbance (e.g., plowing), due to the natural shift in the δ¹³C signature of soil C. For the first time, we used a repeated ¹³C natural abundance approach to measure C turnover in a loamy Gleyic Cambisol after 9 and 21 years of Miscanthus cultivation. This is the longest C₃–C₄ vegetation change study on C turnover in soil under energy crops. SOM stocks under Miscanthus and reference grassland were similar down to 1 m depth. However, both increased between 9 and 21 years from 105 to 140 mg C ha^?1 (P < 0.05), indicating nonsteady state of SOM. This calls for caution when estimating SOM turnover based on a single sampling. The mean residence time (MRT) of old C (>9 years) increased with depth from 19 years (0–10 cm) to 30–152 years (10–50 cm), and remained stable below 50 cm. From 41 literature observations, the average SOM increase after conversion from cropland or grassland to Miscanthus was 6.4 and 0.4 mg C ha^?1, respectively. The MRT of total C in topsoil under Miscanthus remained stable at ~60 years, independent of plantation age, corroborating the idea that C dynamics are dominated by recycling processes rather than by C stabilization. In conclusion, growing Miscanthus on C‐poor arable soils caused immediate C sequestration because of higher C input and decreased SOM decomposition. However, after replacing grasslands with Miscanthus, SOM stocks remained stable and the MRT of old C₃‐C increased strongly with depth.     

Changes in the abundance of C3/C4 species of Inner Mongolia grassland: evidence from isotopic composition of soil and vegetation

Global warming, increasing CO₂ concentration, and environmental disturbances affect grassland communities throughout the world. Here, we report on variations in the C3/C4 pattern of Inner Mongolian grassland derived from soil and vegetation. Soil samples from 149 sites covering an area of approximately 250 000 km² within Inner Mongolia, People's Republic of China were analyzed for the isotopic composition (δ¹³C) of soil organic carbon (SOC). The contrast in δ¹³C between C3 and C4 plants allowed for calculation of the C3/C4 ratio from δ¹³C of SOC with a two‐member mixing model, which accounted for influences of aridity and altitude on δ¹³C of the C3 end‐member and for changes in δ¹³C of atmospheric CO₂. Maps were created geostatistically, and showed a substantially lower C4 abundance in soil than in recent vegetation (?10%). The difference between soil and vegetation varied regionally and was most pronounced within an E–W belt along 44°N and in a mountainous area, suggesting a spread of C4 plants toward northern latitudes (about 1°) and higher altitudes. The areas of high C4 abundance for present vegetation and SOC were well delineated by the isotherms of crossover temperature based on the climatic conditions of the respective time periods. Our study indicates that change in the patterns of C3/C4 composition in the Inner Mongolia grassland was mainly triggered by increasing temperature, which overrode the antagonistic effect of rising CO₂ concentrations.     

Going Beyond Lithium Hybrid Capacitors: Proposing a New High‐Performing Sodium Hybrid Capacitor System for Next‐Generation Hybrid Vehicles Made with Bio‐Inspired Activated Carbon

A novel sodium hybrid capacitor (NHC) is constructed with an intercalation‐type sodium material [carbon coated‐Na₃V₂(PO₄)₃, C‐NVP] and high surface area‐activated carbon derived from an eco‐friendly resource cinnamon sticks (CDCs) in an organic electrolyte. This novel NHC possesses a combination of high energy and high power density, along with remarkable electrochemical stability. In addition, the C‐NVP/CDC system outperforms present, well‐established lithium hybrid capacitor systems in all areas, and can thus be added to the list of candidates for future electric vehicles. A careful optimization of mass balance between electrode materials enables the C‐NVP/CDC cell to exhibit extraordinary capacitance performance. This novel NHC produces an energy density of 118 Wh kg^?1 at a specific power of 95 W kg^?1 and retains an energy density of 60 Wh kg^?1 with high specific power of 850 W kg^?1. Furthermore, a discharge capacitance of 53 F g^?1 is obtained from the C‐NVP/CDC cell at a 1 mA cm^?2 current density, along with 95% capacitance retention, even after 10 000 cycles. The sluggish kinetics of the Na ion battery system is successfully overcome by developing a stable, high‐performing NHC system.     

Leaf economics and hydraulic traits are decoupled in five species‐rich tropical‐subtropical forests

Leaf economics and hydraulic traits are critical to leaf photosynthesis, yet it is debated whether these two sets of traits vary in a fully coordinated manner or there is room for independent variation. Here, we tested the relationship between leaf economics traits, including leaf nitrogen concentration and leaf dry mass per area, and leaf hydraulic traits including stomatal density and vein density in five tropical‐subtropical forests. Surprisingly, these two suites of traits were statistically decoupled. This decoupling suggests that independent trait dimensions exist within a leaf, with leaf economics dimension corresponding to light capture and tissue longevity, and the hydraulic dimension to water‐use and leaf temperature maintenance. Clearly, leaf economics and hydraulic traits can vary independently, thus allowing for more possible plant trait combinations. Compared with a single trait dimension, multiple trait dimensions may better enable species adaptations to multifarious niche dimensions, promote diverse plant strategies and facilitate species coexistence.