Soil heterogeneity buffers community response to climate change in species‐rich grassland

Climate change impacts on vegetation are mediated by soil processes that regulate rhizosphere water balance, nutrient dynamics, and ground‐level temperatures. For ecosystems characterized by high fine‐scale substrate heterogeneity such as grasslands on poorly developed soils, effects of climate change on plant communities may depend on substrate properties that vary at the scale of individuals (<m²), leading to fine‐scale shifts in community structure that may go undetected at larger scales. Here, we show in a long‐running climate experiment in species‐rich limestone grassland in Buxton, England (UK), that the resistance of the community to 15‐year manipulations of temperature and rainfall at the plot scale (9 m²) belies considerable community reorganization at the microsite (100 cm²) scale. In individual models of the abundance of the 25 most common species with respect to climate treatment and microsite soil depth, 13 species exhibited significant soil depth affinities, and nine of these have shifted their position along the depth gradient in response to one or more climate treatments. Estimates of species turnover across the depth gradient reviewed in relation to measurements of water potential, nitrogen supply, pH, and community biomass suggest that communities of shallow microsites are responding directly to microenvironmental changes induced by climate manipulation, while those of the deepest microsites are shifting in response to changes in competitive interference from more nutrient‐demanding species. Moreover, for several species in summer drought and winter heated treatments, climate response in deep microsites was opposite that of shallow microsites, suggesting microsite variation is contributing to community stability at the whole‐plot level. Our study thus demonstrates a strong link between community dynamics and substrate properties, and suggests ecosystems typified by fine‐scale substrate heterogeneity may possess a natural buffering capacity in the face of climate change.     

Positive interactions among plant species for pollinator service: assessing the ‘magnet species’ concept with invasive species

Plants with poorly attractive flowers or with little floral rewards may have inadequate pollinator service, which in turn reduces seed output. However, pollinator service of less attractive species could be enhanced when they are associated with species with highly attractive flowers (so called ‘magnet‐species’). Although several studies have reported the magnet species effect, few of them have evaluated whether this positive interaction result in an enhancement of the seed output for the beneficiary species. Here, we compared pollinator visitation rates and seed output of the invasive annual species Carduus pycnocephalus when grow associated with shrubs of the invasive Lupinus arboreus and when grow alone, and hypothesized that L. arboreus acts as a magnet species for C. pycnocephalus. Results showed that C. pycnocephalus individuals associated with L. arboreus had higher pollinator visitation rates and higher seed output than individuals growing alone. The higher visitation rates of C. pycnocephalus associated to L. arboreus were maintained after accounting for flower density, which consistently supports our hypothesis on the magnet species effect of L. arboreus. Given that both species are invasives, the facilitated pollination and reproduction of C. pycnocephalus by L. arboreus could promote its naturalization in the community, suggesting a synergistic invasional process contributing to an ‘invasional meltdown’. The magnet effect of Lupinus on Carduus found in this study seems to be one the first examples of indirect facilitative interactions via increased pollination among invasive species.     

Individual species–area relationships and spatial patterns of species diversity in a Great Basin,semi‐arid shrubland

Traditional biodiversity metrics operate at the level of a plant community but do not capture spatial variation in diversity from a ‘plant's‐eye view’ of a community. Recently‐developed statistics consider the spatial patterns of plants as well as the number and distribution of species in local plant neighborhoods to quantitatively assess multispecies spatial patterns from a ‘plant's‐eye view’. We used one such statistic, the individual species area relationship (ISAR), to assess spatial patterns of species diversity in a Great Basin (USA) semi‐arid shrubland through an analysis of a spatial dataset on shrub species and locations. In conjunction with appropriate null models, the ISAR blends species area relationships with second‐order spatial statistics to measure the expected species richness in local neighborhoods of variable size around the individuals of a focal species within a community. We found that, contrary to a previous analysis using more traditional methods, the community was well‐mixed with a typical shrub surrounded on average by 4.9 shrub neighbors of 2.1 species at a neighborhood scale of 1.0 m. We also found statistically significant fine‐scale variation in diversity patterns, such that neighborhoods of two species were more diverse than expected by a heterogeneous Poisson null model that accounted for larger‐scale habitat heterogeneity. However, this effect was caused by intraspecific aggregation of these species and was not due to positive interspecific association. Contrary to previous findings in other semi‐arid shrublands, our analysis suggests that the spatial pattern of the shrub community was not significantly structured by interspecific facilitation. This result supports growing evidence for balanced species patterns of adult plants in multispecies communities. Our approach may be used in other communities to describe complex multispecies spatial patterns, quantify species‐specific associations with diversity patterns, and to generate hypotheses regarding relationships between patterns and community‐structuring processes.     

Factors determining distributions of tree species and plant functional types

Plant functional types have been identified by the International Geosphere Biosphere Program as functionally similar basic plant types, especially trees, as needed for global ecological modeling. Based to some extent on an earlier set of pheno-physiognomically defined plant types, a Global Biome Model was produced but has not satisfied all the desired functional criteria posed by IGBP physiologists and modelers. This paper asks two questions: what are the main environmental factors which limit terrestrial plant types, especially tree types; and how many types of potential vegetation are needed to cover the world's terrestrial vegetation patterns? Based on the main environmental factors recognized, a model of world potential dominant vegetation types was produced and used to estimate the minimal number of vegetation types needed. The resulting set of 40 potential dominant vegetation types may serve as an initial basis for a structural-functionally based set of world plant functional types.     

Preference for different inorganic nitrogen forms among plant functional types and species of the Patagonian steppe

We have explored species–specific preferences for nitrate (NO₃ ^?) and ammonium (NH₄ ⁺) as an alternative niche separation in ecosystems where nitrogen (N) is present mostly in inorganic forms. The Patagonian steppe is dominated by shrubs and grasses. Shrubs absorb water and nutrients from deep soil layers, which are poor in N, while grasses have the opposite pattern, absorbing most of their water and nutrients from the upper layers of the soil. We hypothesized that the preferences of shrub and grass for inorganic N forms are different and that the rate of potential N uptake is greater in shrubs than in grasses. To test this hypothesis, we grew individuals of six dominant species in solutions of different NH₄ ⁺:NO₃ ^? concentration ratios. Nitrate uptake was found to be higher in shrubs, while ammonium uptake was similar between plant functional types. The NH₄ ⁺:NO₃ ^? uptake ratio was significantly lower for shrubs than grasses. Shrubs, which under field conditions have deeper rooting systems than grasses, showed a higher N absorption capacity than grasses and a preference for the more mobile N form, nitrate. Grasses, which had lower N uptake rates than shrubs, preferred ammonium over nitrate. These complementary patterns between grasses and shrubs suggest a more thorough exploitation of resources by diverse ecosystems than those dominated by just one functional type. The loss of one plant functional group or a significant change in its abundance would therefore represent a reduction in resource use efficiency and ecosystem functioning.     

The spatial scale of adaptive population differentiation in a wide‐spread,well‐dispersed plant species

Adaptation to the specific conditions at different sites may contribute strongly to the wide distribution of a plant species. However, little is known about the scale at which such adaptation occurs in common species. We studied population differentiation, plasticity and local adaptation of the short‐lived perennial Hypochoeris radicata, a widespread and common plant whose seeds are well‐dispersed. We reciprocally transplanted seedlings among several populations of different size within and among three European regions (in the northwest Czech Republic, central Germany and the central Netherlands) and studied several fitness‐related traits over two growing seasons. The region in which the reciprocal transplant experiment was carried out had no influence on the performance of seedlings, indicating that there were no differences in overall habitat quality. In contrast, the site within region, and the plot within site strongly influenced mean plant performance. Plants from different populations of origin differed in their performance, indicating genetic variation among populations, but performance strongly depended on the specific combination of population of origin and transplant site. Plants that grew at their home site produced on average almost twice the number of seeds per transplant (a multiplicative fitness measure) than foreign plants originating from other sites. Survival, rosette size and multiplicative fitness all decreased with increasing distance from the home site to the transplant site. The size of the population of origin did not influence overall plant performance or the strength of local adaptation. In conclusion, our results indicate that the common and widespread H. radicata consists of locally adapted genotypes within its European range at a relatively small scale. Thus a large potential for gene flow by seeds and a high density of populations do not appear to be sufficient to prevent population differentiation by selection.     

Nutrient limitation in species‐rich Calthion grasslands in relation to opportunities for restoration in a peat meadow landscape

Questions: Which nutrient(s) limit(s) vegetation productivity in Calthion grasslands? Is phosphorus release a bottleneck for restoration of species‐rich Calthion grasslands on rewetted dairy meadows? Location: Three species‐rich Calthion grasslands in the Western Peat District in the Netherlands. Methods: We conducted a field fertilization experiment with nitrogen (N), phosphorus (P) and potassium (K) in three existing Calthion grasslands to evaluate the potential for restoration on rewetted dairy meadows. Responses of above‐ground biomass, tissue nutrient concentrations and nutrient ratios were determined after 2 yr of fertilization. Results: Biomass increased with fertilization with N‐only and K‐only but did not react to P‐only additions. Comparisons of tissue nutrient concentrations and nutrient ratios also gave indications of N and K limitation. Conclusions: The strong P release expected after rewetting should not necessarily interfere with restoration of Calthion communities on rewetted dairy meadows. It is concluded that for successful restoration management measures should focus on reducing N and/or K availability. Potassium might be an overlooked bottleneck in the restoration of species‐rich grasslands.     

Molecular phylogeny of Eois (Lepidoptera,Geometridae): evolution of wing patterns and host plant use in a species‐rich group of Neotropical moths

Strutzenberger, P., Brehm, G., Bodner, F. & Fiedler K. (2010). Molecular phylogeny of Eois (Lepidoptera, Geometridae): evolution of wing patterns and host plant use in a species‐rich group of Neotropical moths. —Zoologica Scripta, 39, 603–620. Eois is a pantropical genus of Geometridae moths with currently 250 valid described species, the majority of which occur in the Neotropics. Eois is a prominent component of Andean moth communities locally accounting for up to ～10% of geometrid individuals. We address the evolution of wing patterns and host plant use in Neotropical Eois and provide a preliminary assessment on the monophyly and biogeographic history of the entire genus as well as affinities within the subfamily Larentiinae. We applied Bayesian, maximum likelihood and maximum parsimony methods of phylogenetic reconstruction to a 142 taxon dataset of partial COI (1220 bp) and Ef1α (1066 bp) sequences resulting in the largest taxon set of geometrid moths analyzed in a molecular phylogenetic study so far. Monophyly of Eois was always strongly supported. Ten monophyletic clades were found with good support, seven of which have characteristic wing pattern phenotypes. Only one wing pattern type occurs in two clades. Trophic associations with representatives of the family Piperaceae occur in all 8 (of 9) Neotropical clades for which host information is available. Apart from feeding on Piper, at least two Eois species in Ecuador feed on Peperomia, and one on Manekia (all Piperaceae); two further species live on Hedyosmum (Chloranthaceae). Species feeding on Peperomia, Manekia and Hedyosmum are usually nested in Piper‐associated clades. Single records of associations with Gesneriaceae and Monimiaceae are scattered in otherwise Piperaceae‐associated clades. These patterns suggest multiple parallel host shifts away from Piper as ancestral food plant. Old World Eois were recovered as monophylum and sister to Neotropical Eois. Within the subfamily Larentiinae the genus Eois has previously been placed close to the tribe Eupitheciini, but this was not supported in our phylogenetic analyses.     

Recovery of soil nitrogen pools in species‐rich grasslands after 12 years of simulated pollutant nitrogen deposition: a 6‐year experimental analysis

Nitrogen (N) deposition from anthropogenic sources is a global problem that can reduce biodiversity and impair ecosystem functioning through effects on soil eutrophication and acidification. While increasing controls on emissions of oxides of nitrogen (NO_x) have reduced European N deposition rates from their peak in the late 20th Century, little is known about the legacy effects of N deposition in soils or the reversibility of N‐induced shifts in ecosystem processes. We studied species‐rich limestone and acidic grasslands, located in a highly polluted region that received over 3000 kg N deposition ha^?1 throughout the 20th Century, followed by a decline of ～50% in NO_x deposition rate in the past two decades. We investigated the effects on seasonal and annual mean concentrations of soil mineral N in experimental plots established in 1990 receiving simulated enhanced N deposition (0–140 kg N ha^?1 yr^?1) until 2002, both in the final year of treatment, and the subsequent 5 years of ‘recovery’ following cessation of treatments. Winter–summer cycles of N mineralization–immobilization were strongly amplified by simulated N deposition rates through the final year of treatments and into the first year of recovery, with winter concentrations of ammonium‐N in the acidic grassland and nitrate in the limestone grassland enhanced by up to 360% and 450%, respectively. Both the magnitude of the seasonal variations and the residual effects of the treatments on soil mineral N concentrations decreased progressively in the first 5 years after treatments ceased, although dose‐dependent trends remained in the acidic grassland. This study establishes that reducing N deposition rates in species‐rich grasslands can reverse eutrophication, even in soils that have experienced prolonged high rates of deposition. It provides new insight into the rates of recovery following, and effects of, declining N deposition rates with implications for restoration of species‐rich grasslands.     

Positive associations between the cushion plant Arenaria polytrichoides (Caryophyllaceae) and other alpine plant species increase with altitude in the Sino‐Himalayas

Question: Does the facilitative effect of cushion plants increase with elevation as a result of increases in environmental harshness? Does this hypothesis apply in the Sino‐Himalayan Mountains? Location: Lakaka Pass on the Baima Snow Mountains (28°20′N, 99°05′E), SW China. Methods: We evaluated the spatial association of several plant species with the cushion plant Arenaria polytrichoides (Caryophyllaceae) at two elevations (4500 m and 4700 m) in the study site and monitored temperature, moisture and nutritional status of soil beneath and outside the cushions. Results: While 14 species grow more frequently associated with the cushions at the higher elevation, at the lower site only three species were positively associated with cushions. Eleven of the species that occurred at both elevations changed their spatial association from neutral or negative with cushions at the lower site to positive at the higher elevation site. Substrate temperatures were rather similar between the cushions and areas of bare ground. Cushions maintained higher moisture than areas of bare ground at both elevations. Soils beneath cushions contained significantly more available nitrogen and potassium compared to open areas at the higher elevation. Conclusions: Our results show that facilitation by A. polytrichoides cushions increases with elevation in the Sino‐Himalayan region. This facilitation effect of A. polytrichoides cushions is probably due to the improved nutrient availability provided by cushion plants in the higher elevation, and these conditions probably permit increased plant recruitment, growth and survival.     

Nestedness and niche‐based species loss in moorland plant communities

Communities in isolated habitat patches surrounded by inhospitable matrices often form a nested subset pattern. However, the underlying causal mechanisms and conservation implications of nestedness in regional communities remain controversial. The nested ranks of species in a nested species‐by‐site matrix may reflect a gradient of species vulnerability to extinction or of colonization ability. However, nestedness analysis has rarely been used to explore determinants of species rank; consequently, little is known of underpinning mechanisms. In this study, we examined nestedness in moorland plant communities widely interspersed within the subalpine zone of northern Japan. Moorland sites differed in area (1000–160 000 m²) and were naturally isolated from one another to various extents within an inhospitable forest matrix. We also determined whether site characteristics (physical and morphometric measures) and species characteristics (niche position and breadth, based on species’ traits) are related to nestedness. Moorland plant communities in the study area were significantly nested. The pH and moorland kernel density (proxy for spatial clustering of moorlands around the focal site) were the most important predictors of moorland site nested rank in a nestedness matrix. Niche breadths of species (measured as variation in leaf mass area and height) predicted species’ nested ranks. Selective environmental tolerances imposed by environmental harshness and selective extinction caused by declines in site carrying capacities probably account for the nested subset pattern in moorland plant communities. The nested rank of species in the nestedness matrix can therefore be translated into the potential order of species loss explainable by species niche breadths (based on variation in functional traits). Complementary understanding of the determinants of site ranking and species ranking in the nestedness matrix provides powerful insight into ecological processes underlying nestedness and into the ways by which communities are assembled or disassembled by such processes.     

Production of small cysteine‐rich effector proteins in Escherichia coli for structural and functional studies

Although the lifestyles and infection strategies of plant pathogens are diverse, a prevailing feature is the use of an arsenal of secreted proteins, known as effectors, which aid in microbial infection. In the case of eukaryotic filamentous pathogens, such as fungi and oomycetes, effector proteins are typically dissimilar, at the protein sequence level, to known protein families and functional domains. Consequently, we currently have a limited understanding of how fungal and oomycete effectors promote disease. Protein biochemistry and structural biology are two methods that can contribute greatly to the understanding of protein function. Both techniques are dependent on obtaining proteins that are pure and functional, and generally require the use of heterologous recombinant protein expression systems. Here, we present a general scheme and methodology for the production and characterization of small cysteine‐rich (SCR) effectors utilizing Escherichia coli expression systems. Using this approach, we successfully produced cysteine‐rich effectors derived from the biotrophic fungal pathogen Melampsora lini and the necrotrophic fungal pathogen Parastagonospora nodorum. Access to functional recombinant proteins facilitated crystallization and functional experiments. These results are discussed in the context of a general workflow that may serve as a template for others interested in understanding the function of SCR effector(s) from their plant pathogen(s) of interest.     

Leaf osmotic potentials of 104 plant species in relation to habitats and plant functional types in Hunshandak Sandland,Inner Mongolia,China

Leaf osmotic potentials ( _s) of 104 plant species from different habitats, i.e., fixed sand dunes, lowland and wetlands in Hunshandak Sandland, Inner Mongolia, China, were investigated. The values of _s were strongly species-specific, and varied from –6.54 MPa ( Caragana microphylla), to –0.44 MPa ( Digitaria ischaemum); 75% of plants investigated had _s from –1.01 to –3.0 MPa. Shrubs were found to have the lowest _s, with an average value of –3.19 MPa, while grasses showed the highest _s. The order of plant _s is shrubs<trees<grasses. The result may relate to anatomical features of shrubs. C₄ photosynthetic pathway plants showed lower _s values. The _s values of 104 species were negatively correlated with their rooting depths ( r ²=0.42; P <0.001). High hydraulic pressure resulting from the deep roots may well explain this trend. The value of _s increased as the environment became wetter, ranging from –0.79 MPa in wetlands to –2.09 MPa in fixed sand dunes. Although soil salt content was higher in wetlands, we did not find any effect on _s.An erratum to this article can be found at     

N‐terminal region of cysteine‐rich protein (CRP) in carlaviruses is involved in the determination of symptom types

Plant viruses in the genus Carlavirus include more than 65 members. Plants infected with carlaviruses exhibit various symptoms, including leaf malformation and plant stunting. Cysteine‐rich protein (CRP) encoded by carlaviruses has been reported to be a pathogenicity determinant. Carlavirus CRPs contain two motifs in their central part: a nuclear localization signal (NLS) and a zinc finger motif (ZF). In addition to these two conserved motifs, carlavirus CRPs possess highly divergent, N‐terminal, 34 amino acid residues with unknown function. In this study, to analyse the role of these distinct domains, we tested six carlavirus CRPs for their RNA silencing suppressor activity, ability to enhance the pathogenicity of a heterologous virus and effects on virus accumulation levels. Although all six tested carlavirus CRPs showed RNA silencing suppressor activity at similar levels, symptoms induced by the Potato virus X (PVX) heterogeneous system exhibited two different patterns: leaf malformation and whole‐plant stunting. The expression of each carlavirus CRP enhanced PVX accumulation levels, which were not correlated with symptom patterns. PVX‐expressing CRP with mutations in either NLS or ZF did not induce symptoms, suggesting that both motifs play critical roles in symptom expression. Further analysis using chimeric CRPs, in which the N‐terminal region was replaced with the corresponding region of another CRP, suggested that the N‐terminal region of carlavirus CRPs determined the exhibited symptom types. The up‐regulation of a plant gene upp‐L, which has been reported in a previous study, was also observed in this study; however, the expression level was not responsible for symptom types.