Facilitation by nurse shrubs of resprouting behavior in a post-fire shrubland in northern Patagonia,Argentina

Abstract. The nurse-plant syndrome is a widely recognized example of positive (facilitative) influences of plant species on the establishment and growth of other species. Most studies of the nurse-plant syndrome have been on species that reproduce mainly from seed rather than vegetatively. In this study, we experimentally compared the influences of two species of nurse shrubs, Schinus patagonicus and Berberis buxifolia, on the survival and growth of vegetatively reproducing herbaceous and woody plants in a post-fire shrubland in northern Patagonia, Argentina. The vegetation beneath shrubs was removed by clipping and, in a paired-sample design, one half of the canopy of each shrub was removed. We determined species richness, counted number of resprouts, and measured photon flux density and soil moisture beneath cut and uncut halves of each shrubs. Abundances of resprouts were several times greater beneath the uncut vs. the cut shrubs, as was the mean number of species. Thus, shrubs have a strong facilitating influence as measured by resprout densities and the number of species. Numbers of resprouts and of species were twice as high beneath Schinus as beneath Berberis, implying important differences in the facilitative effects of the two shrubs species. Microsites beneath Schinus were characterized by lower and more heterogeneous light levels but by greater soil moisture. Even though the reproductive mode in this post-fire shrubland is overwhelmingly vegetative, facilitation by nurse shrubs is important and differentially effective for different species of nurse shrubs.     

Invasion of the alien shrub Prunus laurocerasus in suburban deciduous forests: Effects on native vegetation and soil properties

Most invading alien plants affect native biodiversity and ecosystem functioning. In a field survey, we assessed the impact of the invasive shrub Prunus laurocerasus on the native vegetation and soil properties in suburban deciduous forests in the region of Basel, Switzerland. We installed four pairs of plots in patches of P. laurocerasus and in adjacent not invaded areas in each of twelve forest areas. Native species richness, Shannon-diversity and species composition of the ground vegetation and shrub layer were assessed in each plot. Furthermore, in each plot we measured physical and soil chemical characteristics, enzyme activities and the carbon source utilization pattern of the soil microbial community using Ecoplates™. The maximum age of P. laurocerasus in each plot was determined using tree ring analysis, indicating the time elapsed since the invasive plant has established. A lower native plant species richness in both the ground vegetation and shrub layer was observed in plots with presence of P. laurocerasus. A different species composition of the ground vegetation was also found among plots with and without the invasive shrub. Plots invaded by P. laurocerasus had a lower soil moisture content than control plots. The intensity, diversity and substrate richness of the carbon sources were increased in soil from invaded plots compared to soil in control plots. However, the chemical soil characteristics examined and the activities of enzymes were not influenced by the invasive plant. The effects of P. laurocerasus became more pronounced with the time elapsed since the invasive plant has established. Thus, the removal of young P. laurocerasus individuals would be an appropriate management practice for this invasive shrub species.     

Patterns of species diversity and soil nutrients along a chronosequence of vegetation recovery in Hainan Island, South China

Understanding the pattern of species diversity and soil factors can enhance our knowledge of the mechanism of vegetation recovery, however, there is still a gap in the knowledge of succession rate and trend for species diversity in relation to soil nutrients during the vegetation recovery process. Patterns of species diversity and soil nutrients during the tropical vegetation recovery as well as the correlation between species diversity and soil nutrients were explored in Hainan Island, located in southern China. Plots assigned as grassland stage (GS), shrub stage (SS), secondary forest stage (SFS), and primary forest stage (PFS) were established using a chronosequence approach. Results showed that species richness and evenness increased from GS to PFS. Species dominance/diversity curves were fitted using the lognormal distribution model (r ² ?=?0.891?C0.972). Species richness for the herb layer was maximal at SFS, whereas species richness for both the shrub layer and tree layer reached their maximum at PFS. Species turnover and soil total phosphorus decreased, whereas organic matter and total nitrogen increased from GS to PFS. Organic matter and total nitrogen were both positively correlated with species richness and total coverage, and total phosphorus was positively correlated with species turnover. The results clearly demonstrate that diversity asymptotically increases and positively correlates with increasing soil fertility, and the total phosphorus value is predicted to be an important soil factor that affects successional rate during tropical vegetation recovery processes.     

Altered N,P and C dynamics with absence of fire in Eucalyptus forests affected by premature decline

Absence of fire is increasingly recognized as an important driver of soil nutrient budgets in Eucalyptus forest, especially in forests affected by premature Eucalyptus decline, due to the effects of soil nutrient accumulation on nutrient balances and forest community dynamics. In this study, we present a dataset of soil and foliar nutrient analyses, and vegetation measurements from a fire chronosequence survey in native E. delegatensis forest. Measured indices include total soil and extractable soil nitrogen (N), or phosphorus (P), soil organic carbon (C), soil acid‐phosphatase (PME) activity, foliar N and foliar P, and understorey and overstorey vegetation canopy height. We show that in some cases indices are strongly linked to time since fire (2–46 years). Time since fire correlated positively with foliar N, total and extractable soil N, soil organic C, and also soil PME activity; the latter an indicator of biotic P demand. Differences in the strength of these relationships were apparent between two geology types, with stronger relationships on the potentially less‐fertile geology. The strong positive correlation with time since fire and understorey canopy height reflected increasing shrub biomass and thickening of the shrub layer. The strong positive correlation for soil or foliar N, but not P, with time since fire, indicates that P does not increase relative to N over time. P may, therefore, become limiting to growth in this plant community. Similarly, the significantly higher concentrations of soil N but not P, also found in both older and long‐unburnt forest stands (>100 years since management), may exacerbate a situation of soil nutrient limitation over several decades. A characteristic feature of long unmanaged stands is a developing tea tree (Leptospermum sp.) understorey, which may benefit from elevated soil N availability and increasing organic C accumulation with prolonged fire absence. This increased shrub biomass would outcompete Eucalyptus for resources, including soil nutrients and water.     

Afforestation causes changes in post‐fire regeneration in native shrubland communities of northwestern Patagonia,Argentina

Question: What are the effects of fire in native shrubland communities and in pine plantations established in these shrublands? Location: Northern Patagonia, Argentina. Methods: We surveyed four sites in Chall‐Huaco valley, located in northwest Patagonia. Each site was a vegetation mosaic composed of an unburned Pinus ponderosa plantation, a plantation burned in 1996, and an unburned matorral and a matorral burned by the same fire. We recorded the cover of all vascular plant species. We also analysed species richness, total cover, proportion of exotic species, abundance of woody species and herb species, cover of exotic species, abundance of woody and herb species and differences in composition of species. For both shrubs and tree species we recorded the main strategy of regeneration (by resprouting or by seed). Results: We found that fire had different effects on native matorral and pine plantations. Five years after fire, plantations came to be dominated by herbs and exotic species, showing differences in floristic composition. In contrast, matorral communities remained very similar to unburned matorral in terms of species richness, proportion of woody species, and herb species and proportion of exotics. Also, pine plantations were primarily colonized by seedlings, while matorrals were primarily colonized by resprouting. Conclusions: Matorrals are highly fire resilient communities, and the practice of establishing plantations on matorrals produces a strong reduction in the capacity of matorral to return to its original state. The elimination of shrubs owing to the effect of plantations can hinder regeneration of native ecosystems. Burned plantations may slowly develop into ecosystems similar to the native ones, or they may produce a new ecosystem dominated by exotic herbs. This study shows that plantations of exotic conifers affect native vegetation even after they have been removed, as in this case by fire.     

Nitrogen-addition effects on leaf traits and photosynthetic carbon gain of boreal forest understory shrubs

Boreal coniferous forests are characterized by fairly open canopies where understory vegetation is an important component of ecosystem C and N cycling. We used an ecophysiological approach to study the effects of N additions on uptake and partitioning of C and N in two dominant understory shrubs: deciduous Vaccinium myrtillus in a Picea abies stand and evergreen Vaccinium vitis-idaea in a Pinus sylvestris stand in northern Sweden. N was added to these stands for 16 and 8 years, respectively, at rates of 0, 12.5, and 50 kg N ha^?1 year^?1. N addition at the highest rate increased foliar N and chlorophyll concentrations in both understory species. Canopy cover of P. abies also increased, decreasing light availability and leaf mass per area of V. myrtillus. Among leaves of either shrub, foliar N content did not explain variation in light-saturated CO₂ exchange rates. Instead photosynthetic capacity varied with stomatal conductance possibly reflecting plant hydraulic properties and within-site variation in water availability. Moreover, likely due to increased shading under P. abies and due to water limitations in the sandy soil under P. sylvestris, individuals of the two shrubs did not increase their biomass or shift their allocation between above- and belowground parts in response to N additions. Altogether, our results indicate that the understory shrubs in these systems show little response to N additions in terms of photosynthetic physiology or growth and that changes in their performance are mostly associated with responses of the tree canopy.     

Woody encroachment reduces nutrient limitation and promotes soil carbon sequestration

During the past century, the biomass of woody species has increased in many grassland and savanna ecosystems. As many of these species fix nitrogen symbiotically, they may alter not only soil nitrogen (N) conditions but also those of phosphorus (P). We studied the N‐fixing shrub Dichrostachys cinerea in a mesic savanna in Zambia, quantifying its effects upon pools of soil N, P, and carbon (C), and availabilities of N and P. We also evaluated whether these effects induced feedbacks upon the growth of understory vegetation and encroaching shrubs. Dichrostachys cinerea shrubs increased total N and P pools, as well as resin‐adsorbed N and soil extractable P in the top 10‐cm soil. Shrubs and understory grasses differed in their foliar N and P concentrations along gradients of increasing encroachment, suggesting that they obtained these nutrients in different ways. Thus, grasses probably obtained them mainly from the surface upper soil layers, whereas the shrubs may acquire N through symbiotic fixation and probably obtain some of their P from deeper soil layers. The storage of soil C increased significantly under D. cinerea and was apparently not limited by shortages of either N or P. We conclude that the shrub D. cinerea does not create a negative feedback loop by inducing P‐limiting conditions, probably because it can obtain P from deeper soil layers. Furthermore, C sequestration is not limited by a shortage of N, so that mesic savanna encroached by this species could represent a C sink for several decades.     

Rapid development of phosphorus limitation in temperate rainforest along the Franz Josef soil chronosequence

The aim of this study was to examine how shifts in soil nutrient availability along a soil chronosequence affected temperate rainforest vegetation. Soil nutrient availability, woody plant diversity, composition and structure, and woody species leaf and litter nutrient concentrations were quantified along the sequence through ecosystem progression and retrogression. In this super-wet, high leaching environment, the chronosequence exhibited rapid soil development and decline within 120,000 years. There were strong gradients of soil pH, N, P and C, and these had a profound effect on vegetation. N:P_leaf increased along the chronosequence as vegetation shifted from being N- to P- limited. However, high N:P_leaf ratios, which indicate P-limitation, were obtained on soils with both high and low soil P availability. This was because the high N-inputs from an N-fixing shrub caused vegetation to be P-limited in spite of high soil P availability. Woody species nutrient resorption increased with site age, as availability of N and P declined. Soil P declined 8-fold along the sequence and P resorption proficiency decreased from 0.07 to 0.01%, correspondingly. N resorption proficiency decreased from 1.54 to 0.26%, corresponding to shifts in mineralisable N. Woody plant species richness, vegetation cover and tree height increased through ecosystem progression and then declined. During retrogression, the forest became shorter, more open and less diverse, and there were compositional shifts towards stress-tolerant species. Conifers (of the Podocarpaceae) were the only group to increase in richness along the sequence. Conifers maintained a lower N:P_leaf than other groups, suggesting superior acquisition of P on poor soils. In conclusion, there was evidence that P limitation and retrogressive forests developed on old soils, but N limitation on very young soils was not apparent because of inputs from an abundant N-fixing shrub.Electronic Supplementary Material Supplementary material is available in the online version of this article at .     

Importance of the soil seed bank of disturbed sites in Chilean matorral in early secondary succession

Abstract. The dynamics of the seed bank may provide clues to the process of recovery of the vegetation of disturbed sites. The role of the seed bank may be more important in areas with a seasonal climate than in areas where seedling recruitment is not limited to one season. We studied the seed bank and the seed rain in three sites of the Chilean mediterranean-climate region (33° 48'S) which differed in the degree of anthropic disturbance: a closed-canopy, second-growth forest; an open matorral; and an old-field. Additionally, we tested the germination of seeds from the soil and from the current-year seed crop. The seed bank varied considerably between the two years of study, although no change in the vegetation could be observed. Seed density and species richness were lower in 1989 than in 1988. The seed bank of the second-growth forest changed less between years, while the old-field showed the largest change. The highest seed rain occurred under shrub patches in the open matorral, while few seeds fell in the spaces between shrub clumps or in the old-field. In the forest, seed rain was low and correlated with species cover. Germination was low (0 - 15%) in tests using either soil samples or fresh seeds. These results indicate that matorral succession is a very slow process, limited mainly by low germination and low arrival of propagules to open areas. Most woody species have animal-disseminated fleshy propagules. The presence of established shrubs which may serve as perches or refuges for animals increases species richness in the seed rain and the seed bank.     

15N fractionation between vegetation, soil, faeces and wool is not influenced by stocking rate

Understanding stable isotope fractionation in trophic networks is important for the interpretation of stable isotope composition of ecosystem components. This work explores the influence of grazing pressure on the nitrogen isotope composition (?? ¹⁵N) of vegetation (standing biomass), soil, and sheep??s faeces and wool in a three-years (2005?C2007) experiment with different stocking rates (0.375?C2.25 sheep ha^-1 year^-1) in semi-arid Inner Mongolia grassland. The ¹⁵N of wool (from a yearly shearing) reflects vegetation at the whole-year grazing grounds-scale while faeces reflect that of the area grazed within a few days. Stocking rate had no effect on ?? ¹⁵N of vegetation and soil, and sheep??s faeces and wool, although nitrogen content of bulk vegetation increased with stocking rate. Furthermore, ?? ¹⁵N of vegetation and diet did not differ between stocking rates. Hence, ¹⁵N fractionations between vegetation and faeces (??_veg-faeces), vegetation and wool (?? _veg-wool), faeces and soil (?? _faeces-soil) and soil and vegetation (?? _soil-veg) were constants, with ?? _veg-faeces?=?3.0?? (±0.1??, 95% confidence interval), ?? _veg-wool?=?5.3?? (±0.1??), ?? _faeces-soil?=?1.1?? (±0.4??) and ?? _soil-veg?=?-4.1?? (±0.3??). This finding is useful as it means that ?? ¹⁵N of wool or faeces can be used to estimate the ¹⁵N of grazed vegetation, even if grazing pressure is unknown.     

Soil fertility and herb facilitation mediated by Retama sphaerocarpa

Abstract. Woody legumes growing in dry climates can increase soil nutrient content and facilitate plant growth in their understorey. We investigated differences in soil fertility and herbaceous community in relation to the presence and absence of the legume Retama sphaerocarpa in a shrubland in a mediterranean type climate. The results showed a higher content of limiting nutrients for plant growth, such as phosphorus and nitrogen, under the R. sphaerocarpa canopy. Herbaceous biomass, cover and nitrogen content increased below the canopy. However, species richness and diversity were diminished in the presence of a dense canopy of R. sphaerocarpa. Nitrogen isotopic analysis did not indicate a clear relationship between symbiotic fixation in R. sphaerocarpa and nitrogen content of soil and plants under its canopy. Nevertheless, herbs growing in the understorey showed a linear correlation between foliar N content and ¹⁵N values. The existence of a dense shrub canopy induced a smaller monthly variation in herb richness, diversity and biomass, suggesting that it provides a stable microhabitat facilitating herbaceous vegetation establishment and growth. The ‘shelter’ effect was more important when environmental conditions became stressful for herbaceous plants.     

Leaf traits in Chilean matorral: sclerophylly within,among, and beyond matorral,and its environmental determinants

Studies of leaf traits often focus on tradeoffs between growth and resource conservation, but little is known about variation in the mechanical traits that influence resource conservation. This study investigates how leaf mechanical traits vary across matorral vegetation in central Chile, how they correlate with environmental factors, and how these trends compare at a broader geographic scale. Leaf toughness, strength, stiffness, and associated traits were measured in five matorral types in central Chile, and relationships with soil N and P and climate variables were assessed. Trends with soil and climate were then analyzed across shrubland and woodland in Chile, Western Australia, and New Caledonia. Chilean species varied in leaf mechanics and associated traits, both within and among matorral types, with more species in sclerophyll matorral having strong, tough, and stiff leaves than in arid and littoral matorral. Overall, leaves with high leaf dry mass per area were stiffer, tougher, stronger, thicker, denser, with more fiber, lignin, phenolics and fiber per unit protein and less protein: tannin activity and N and P per mass, forming a broad sclerophylly syndrome. Mechanical traits of matorral species were not correlated with soil N or P, or predictably with climate variables, except flexural stiffness (EI_W) which correlated positively with annual reference evapotranspiration (ET₀). However, soil P made strong independent contributions to variation in leaf mechanics across shrublands and woodlands of Chile, Western Australia, and New Caledonia, either separately (strength) or together with ET₀ (toughness) explaining 46–90% of variation. Hence ET₀ was predictive of EI_W in Chilean matorral, whereas soil P was highly predictive of variation in leaf strength, and combined with ET₀ was highly predictive of toughness, at a broader geographic scale. The biological basis of these relationships, however, may be complex.     

Factors determining plant species richness in Alaskan arctic tundra

Abstract. We studied the relationship between plant N:P ratio, soil characteristics and species richness in wet sedge and tussock tundra in northern Alaska at seven sites. We also collected data on soil characteristics, above‐ground biomass, species richness and composition. The N:P ratio of the vegetation did not show any relationship with species richness. The N:P ratio of the soil was related with species richness for both vegetation types. Species richness in the tussock tundra was most strongly correlated with soil calcium content and soil pH, with a strong correlation between these two factors. N:P ratio of the soil was also correlated with soil pH. Other factors correlated with species richness were soil moisture and Sphagnum cover. Organic matter content was the factor most strongly correlated with species richness in the wet sedge vegetation. N:P ratio of the soil was strongly correlated with organic matter content. We conclude that N:P ratio in the vegetation is not an important factor determining species richness in arctic tundra and that species richness in arctic tundra is mainly determined by pH and flooding. In tussock tundra the pH, declining with soil age, in combination with Sphagnum growth strongly decreases species richness, while in wet sedge communities flooding over long periods of time creates less favourable conditions for species richness.     

Riparian zones as havens for exotic plant species in the central grasslands

In the Central Grasslands of the United States, we hypothesized that riparian zones high in soil fertility would contain more exotic plant species than upland areas of low soil fertility. Our alternate hypothesis was that riparian zones high in native plant species richness and cover would monopolize available resources and resist invasion by exotic species. We gathered nested-scale vegetation data from 40 1 m²subplots (nested in four 1000 m² plots) in both riparian and upland sites at four study areas in Colorado, Wyoming, and South Dakota (a total of 320 1 m² subplots and 32 1000 m² plots). At the 1 m² scale, mean foliar cover of native species was significantly greater (P<0.001) in riparian zones (36.3% ± 1.7%) compared to upland sites (28.7% ± 1.5%), but at this small scale there were no consistent patterns of native and exotic species richness among the four management areas. Mean exotic species cover was slightly higher in upland sites compared to riparian sites (9.0% ± 3.8% versus 8.2% ± 3.0% cover). However, mean exotic species richness and cover were greater in the riparian zones than upland sites in three of four management areas. At the 1000 m² scale, mean exotic species richness was also significantly greater (P<0.05) in riparian zones (7.8 ± 1.0 species) compared to upland sites (4.8 ± 1.0 species) despite the heavy invasion of one upland site. For all 32 plots combined, 21% of the variance in exotic species richness was explained by positive relationships with soil % silt (t =1.7, P=0.09) and total foliar cover (t = 2.4, P=0.02). Likewise, 26% of the variance in exotic species cover (log₁₀ cover) was explained by positive relationships with soil % silt (t =2.3, P=0.03) and total plant species richness (t = 2.5, P=0.02). At landscape scales (four 1000 m² plots per type combined), total foliar cover was significantly and positively correlated with exotic species richness (r=0.73, P<0.05) and cover (r=0.74, P<0.05). Exotic species cover (log₁₀ cover) was positively correlated with log₁₀% N in the soil (r=0.61, P=0.11) at landscape scales. On average, we found that 85% (±5%) of the total number of exotic species in the sampling plots of a given management area could be found in riparian zones, while only 50% (±8%) were found in upland plots. We conclude that: (1) species-rich and productive riparian zones are particularly invasible in grassland ecosystems; and (2) riparian zones may act as havens, corridors, and sources of exotic plant invasions for upland sites and pose a significant challenge to land managers and conservation biologists.     

Variation of 15N natural abundance in leaves and nodules of actinorhizal shrubs in Northwest Patagonia

This study was performed to assess the N₂-fixing capability of the native actinorhizal species Ochetophila trinervis (sin. Discaria trinervis) and Discaria chacaye (Rhamnaceae) in Northwest Patagonia. We measured the N concentration and ¹⁵N natural abundance in leaves and nodules of O. trinervis and D. chacaye, in leaves of associated non-actinorhizal vegetation, and in the soils under each sampled plant. O. trinervis and D. chacaye had foliar N concentrations that were about twice that of non-actinorhizal shrubs growing at the same sites, even though soils varied four-fold in total N across the sites. Leaves of both actinorhizal plants had a similar δ¹⁵N at any site and were close to atmospheric values. The foliar δ¹⁵N of non-actinorhizal plants and soil δ¹⁵N were strongly correlated across the sites. Nodules were depleted in δ¹⁵N relative to the foliage of the respective actinorhizal species. In conjunction with the uniformly high foliage N concentration of these actinorhizal plants and the universal presence of vesicles observed in root nodules, these data strongly suggest that O. trinervis and D. chacaye obtain a significant amount of their N from N₂ fixation. To calculate the proportion of N derived from atmosphere, theoretical B-values were estimated. In all cases where the δ¹⁵N of fixing and reference foliage were significantly different, O. trinervis and D. chacaye obtained almost all of their N from N₂ fixation. These results are the first to demonstrate N₂ fixation by O. trinervis and D. chacaye in the field and therefore suggest an important role for these actinorhizal plants in the N economy of ecosystems in northwest Patagonia as well as their potential use for restoration of degraded lands in this region.     

荒漠草原区柠条固沙人工林地表草本植被季节变化特征

研究荒漠草原人工林固沙区地表草本植被季节变化特征及其和柠条林龄的关系,对于分析柠条人工林地表草本植物的季节适应性和制订合理的人工林管理措施均具有重要的科学意义。选择6、15、24年生和36年生柠条人工林为研究对象,通过调查每个样地5月、8月和10月地表草本植物密度、物种数、盖度和高度,分析了荒漠草原区柠条人工固沙林生长过程中地表草本植被季节变化特征及其影响因素。结果表明,地表草本植物物种数在柠条林龄6和15a时受季节改变的影响较小,在24a之后受到季节变化的显著影响(P0.05)。地表草本植物密度在柠条林龄6a时受季节改变的影响较小,但在15a之后季节变化显著影响地表草本植物个体数分布(P0.05),而且在10月具有最多的地表草本植物个体数。地表草本植被盖度和高度均受到季节变化的显著影响(P0.05),而受林龄的影响较小;不同年龄林地地表草本植被盖度和高度均表现为10月和8月较高,5月较低。研究表明,荒漠草原柠条人工林固沙区,柠条林发育生长和灌木形态的改变不仅影响土壤营养条件,而且还可以调控由于季节改变而引起的土壤温湿度变化,柠条林龄和季节更替二者交互作用,共同影响地表草本植被的季节变化特征。     

Canopy effects of the invasive shrub Pyracantha angustifolia on seed bank composition,richness and density in a montane shrubland (Córdoba,Argentina)

Abstract Invasive woody species frequently change the composition of the established vegetation and the properties of the soil under their canopies. Accordingly, invasion may well affect regenerative phases of the community, especially at the seed bank level, likely influencing community restoration. Pyracantha angustifolia (Rosaceae) is an invasive shrub in central Argentina that affects woody recruitment, particularly enhancing the recruitment of other exotic woody species. There is though no information regarding its effect on the soil seed bank within the invaded community. The present study was set up to gain further insight into the canopy effects of P. angustifolia. We aimed to assess whether the invasive shrub affects seed bank composition, richness and seed density as compared with the dominant native shrub Condalia montana (Rhamnaceae), and to relate the observed seed bank patterns with those of the established vegetation. We evaluated the composition of the germinable seed bank and the established vegetation under the canopy of 16 shrubs of P. angustifolia, 16 shrubs of C. montana, and in 16 control plots (10 m²) without shrub cover. The floristic composition of the seed bank differed among canopy treatments. However, seed bank richness did not differ significantly. There was an overall high seed density of exotic species throughout the study site, though exotic forbs showed significantly lower seed densities under the invasive shrub. Pyracantha angustifolia would not promote the incorporation of new species into the seed bank of the invaded community but rather favour the establishment of woody species that do not depend on seed banks. The absence of dominant woody species in the seed bank, the dominance of exotic forbs, and the high similarity between established exotic species and those present in the seed bank may surely affect community restoration following the main disturbances events observed in the region.     

Linking water use and nutrient accumulation in tree island upland hammock plant communities in the Everglades National Park, USA

The tree island hammock communities in the Florida Everglades provide one of many examples of self-organized wetland landscape. However, little is understood about why these elevated tree island communities have higher nutrient concentration than the surrounding freshwater marshes. Here we used stable isotopes and elemental analysis to compare dry season water limitation and soil and foliar nutrient status in upland hammock communities of 18 different tree islands located in the Shark River Slough and adjacent prairie landscapes. We observed that prairie tree islands, having a shorter hydroperiod, suffer greater water deficits during the dry season than slough tree islands by examining shifts in foliar ??¹³C values. We also found that prairie tree islands have lower soil total phosphorus concentration and higher foliar N/P ratio than slough tree islands. Foliar ??¹⁵N values, which often increase with greater P availability, was also found to be lower in prairie tree islands than in slough tree islands. Both the elemental N and P and foliar ??¹⁵N results indicate that the upland hammock plant communities in slough tree islands have higher amount of P available than those in prairie tree islands. Our findings are consistent with the transpiration driven nutrient harvesting chemohydrodynamic model. The water limited prairie tree islands hypothetically transpire less and harvest less P from the surrounding marshes than slough tree islands during the dry season. These findings suggest that hydroperiod is important to nutrient accumulation of tree island habitats.     

Belowground competitive suppression of seedling growth by grass in an African savanna

Coexistence of N₂-fixing legumes and non-legume trees with grasses in African savannas results in intense competition between these life-forms. We hypothesised that belowground competition might induce different nutritional constraints in N₂- versus non-N₂-fixing species. A field (Hluhluwe-imFolozi nature reserve, South Africa) competition experiment with two N₂-fixing legume species (Acacia burkei and Acacia karroo) and two non-N₂-fixing species (Schotia brachypetala and Spirostachys africana) both with clipped grass and without grass was established. Plants were supplied with no fertilizer, or generous amounts of fertilizer (200?kg?N?ha^?1, 100?kg P₂O₅ ha^?1, 7.1?kg K₂O ha^?1) supplied as either 28?C10 (N?CK), P or a combination of these fertilizers (NPK). Regularly clipped grass suppressed growth (by more than 90?%) of both N₂- and non-N₂-fixing seedlings equally. Biomass accumulation of seedlings grown with grass and the grasses themselves responded positively to NK and/or NPK, but not P, although P-fertilization did have effects on foliar [N] and ??¹⁵N values of trees and grasses showing that plants accessed the fertilizer. Tree ??¹⁵N values and foliar [N] were also modified by NPK, demonstrating access to fertilizer. However, the ameliorative effects of NPK on grass competition-induced biomass suppression were only partial. This may be due to ??non-resource competition?? (i.e. root gaps) imposed by dense grass roots. The fact that nutrients were able to partially ameliorate the effects of grass competition, however, indicates that such ??non-resource competition?? may be partially overcome by even more generous supply of fertilizers.     

Nitrogen fertilization and fire act independently on foliar stoichiometry in a temperate steppe

Nitrogen (N) fertilization, as a grassland management strategy, has been widely used to improve forage quality and increase the productivity of grasslands degraded by overstocking. It is widely accepted that N addition will alter ecosystem structure and function, and that these effects may be altered by natural disturbances, such as fire. We examined the effects of annual burning and N fertilization (17.5 g N m^?2 year^?1, at a surplus rate in order to simulate agriculture treatment) on foliar chemistry and stoichiometric ratios of eight dominant species (Leymus chinensis, Stipa grandis, Cleistogenes squarrosa, Potentilla bifurca, Thalictrum squarrosum, Artemisia frigida, Kochia prostrata and Caragana microphylla) in a temperate steppe in northern China. After 3 years of treatments, annual burning significantly increased soil extractable phosphorus (P) concentration but showed no effects on soil inorganic N concentration, whereas N fertilization caused a significant increase in inorganic N concentration but not of extractable P. Species differed substantially with respect to all nutritional and stoichiometric variables. Both annual burning and N fertilization caused significant increases in foliar N and P concentrations and thus decreases in carbon (C):N and C:P ratios. While annual burning showed no effects on N:P ratios, N fertilization produced higher N:P ratios. However, species responded idiosyncratically to both fire and N fertilization in terms of foliar N concentration, C:N and N:P ratio. In addition, there was no interaction between fire and N fertilization that affected all variables. This study suggests that both annual burning and N fertilization have direct impacts on plant elemental composition and that fire- and N addition-induced changes of community composition may have important consequences for plant-mediated biogeochemical cycling pathways in temperate steppe ecosystem.