Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra‐ and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five‐fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra‐ and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.     

Synchronous fire activity in the tropical high Andes: an indication of regional climate forcing

Global climate models suggest enhanced warming of the tropical mid and upper troposphere, with larger temperature rise rates at higher elevations. Changes in fire activity are amongst the most significant ecological consequences of rising temperatures and changing hydrological properties in mountainous ecosystems, and there is a global evidence of increased fire activity with elevation. Whilst fire research has become popular in the tropical lowlands, much less is known of the tropical high Andean region (>2000masl, from Colombia to Bolivia). This study examines fire trends in the high Andes for three ecosystems, the Puna, the Paramo and the Yungas, for the period 1982–2006. We pose three questions: (i) is there an increased fire response with elevation? (ii) does the El Niño‐ Southern Oscillation control fire activity in this region? (iii) are the observed fire trends human driven (e.g., human practices and their effects on fuel build‐up) or climate driven? We did not find evidence of increased fire activity with elevation but, instead, a quasicyclic and synchronous fire response in Ecuador, Peru and Bolivia, suggesting the influence of high‐frequency climate forcing on fire responses on a subcontinental scale, in the high Andes. ENSO variability did not show a significant relation to fire activity for these three countries, partly because ENSO variability did not significantly relate to precipitation extremes, although it strongly did to temperature extremes. Whilst ENSO did not individually lead the observed regional fire trends, our results suggest a climate influence on fire activity, mainly through a sawtooth pattern of precipitation (increased rainfall before fire‐peak seasons (t‐1) followed by drought spells and unusual low temperatures (t0), which is particularly common where fire is carried by low fuel loads (e.g., grasslands and fine fuel). This climatic sawtooth appeared as the main driver of fire trends, above local human influences and fuel build‐up cyclicity.     

Directed dispersal by rotational shepherding supports landscape genetic connectivity in a calcareous grassland plant

Directed dispersal by animal vectors has been found to have large effects on the structure and dynamics of plant populations adapted to frugivory. Yet, empirical data are lacking on the potential of directed dispersal by rotational grazing of domestic animals to mediate gene flow across the landscape. Here, we investigated the potential effect of large‐flock shepherding on landscape‐scale genetic structure in the calcareous grassland plant Dianthus carthusianorum, whose seeds lack morphological adaptations to dispersal to animals or wind. We found a significant pattern of genetic structure differentiating population within grazed patches of three nonoverlapping shepherding systems and populations of ungrazed patches. Among ungrazed patches, we found a strong and significant effect of isolation by distance (r = 0.56). In contrast, genetic distance between grazed patches within the same herding system was unrelated to geographical distance but significantly related to distance along shepherding routes (r = 0.44). This latter effect of connectivity along shepherding routes suggests that gene flow is spatially restricted occurring mostly between adjacent populations. While this study used nuclear markers that integrate gene flow by pollen and seed, the significant difference in the genetic structure between ungrazed patches and patches connected by large‐flock shepherding indicates the potential of directed seed dispersal by sheep across the landscape.     

Directional climate change and potential reversal of desertification in arid and semiarid ecosystems

Our objective was to determine if long‐term increases in precipitation can maintain grasslands susceptible to desertification, and initiate a reversal of historic regime shifts on desertified shrublands. Perennial grass production and species richness in a multi‐year wet period were hypothesized to be greater than expected based on precipitation in a sequence of dry years. These responses were expected to differ for grasslands and shrublands with different dominant species and topo‐edaphic properties. Long‐term trends in desertification were documented using vegetation maps beginning in 1858, 1915, 1928, and 1998). These trends were compared with herbaceous and woody species responses to a sequence of dry (1994–2003) and wet years (2004–2008) for two grassland (uplands, playas) and three desertified shrubland types (honey mesquite, creosotebush, tarbush) in the Chihuahuan Desert. Analyses showed that both types of grasslands decreased in spatial extent since 1858 whereas areas dominated by mesquite or creosotebush increased. Production of upland grasslands in the wet period was greater than expected based on responses during the dry period whereas the relationships between species richness and precipitation was the same for both periods. Precipitation was not important to responses in playa grasslands in either period. For all ecosystem types, the production response in wet years primarily was an increase in herbaceous plants, and the most pronounced responses occurred on sandy sites (upland grasslands, mesquite shrubland). Results suggest that multiple wet years are needed to initiate a sequence of grass establishment and survival processes that can maintain upland grasslands without management inputs and lead to a state change reversal in desertified shrublands. Restoration strategies need to take advantage of opportunities provided by future climates while recognizing the importance of ecosystem type.     

Effect of geographical range size on plant functional traits and the relationships between plant,soil and climate in Chinese grasslands

Aim Our aim was to address the potential effect of the geographical range size of species on the relationships between plant traits, soil and climate in Chinese grasslands. Previous analyses tended to examine plant–environment relationships across many species while ignoring that species with different range sizes may respond differently to the environment. Here we hypothesized that leaf traits of narrow‐ranging species would be more strongly correlated with soil and climatic variables than those of wide‐ranging species. Location Chinese grasslands. Methods Data on leaf traits, including nitrogen and phosphorus concentrations, carbon/nitrogen ratio, nitrogen/phosphorus ratio and specific leaf area, as well as species range sizes for 208 species distributed across 178 sites in Chinese grasslands were collected. Soil and climate information for each study site was also gathered. The effects of range size on leaf traits were tested using one‐way ANOVA. Correlations between leaf traits, soil and climate were calculated for all species pooled together and for species partitioned into range size quartiles, from the first (narrowest‐ ranging 25%) to the fourth (widest‐ranging 25%). Results Narrow‐ranging species tended to occur at high altitude with lower temperature but higher soil nutrient concentrations compared with wide‐ranging species. No direct link between leaf traits and species range sizes was detected. However, patterns of leaf–soil nutrient relationships changed significantly across levels of range size. Narrow‐ranging species tended to be more sensitive to variation in soil nutrient availability than wide‐ranging species, resulting in a shift from a positive leaf–soil nutrient relationship for narrow‐ranging plants to no relationship for wide‐ranging plants. Species responses to climatic variables were unrelated to their range sizes. Main conclusions The close relationship between leaf and soil nutrients indicates a specialization of narrow‐ranging species to particular habitats whereas wide‐ranging species may be able to better withstand changes in environment such as soil fertility over a large area.     

Nutrient concentration ratios and co-limitation in South African grasslands

*Assessing plant nutrient limitation is a fundamental part of understanding grassland dynamics. The ratio of concentrations of nitrogen (N) and phosphorus (P) in vegetation has been proposed as an index of the relative limitation of biomass production by N and P, but its utility has not been tested well in grasslands. *At five sites in Kruger National Park, South Africa, across soil and precipitation contrasts, N and P were added in a factorial design to grass-dominated plots. *Although the N:P ratio of unfertilized vegetation across all sites (5.8) would have indicated that production was N-limited, aboveground production was consistently co-limited by N and P. Aboveground production was still greater in plots fertilized with N and P than in those fertilized with just N, but the N:P ratio did not exceed standard thresholds for P limitation in N-fertilized vegetation. Comparisons among sites showed little pattern between site N:P ratio and relative responses to N and P. *When combined with results from other grassland fertilization studies, these data suggest that the N:P ratio of grasses has little ability to predict limitation in upland grasslands. Co-limitation between N and P appears to be much more widespread than would be predicted from simple assumptions of vegetative N:P ratios.     

Net ecosystem carbon exchange in two experimental grassland ecosystems

Increases in net primary production (NPP) may not necessarily result in increased C sequestration since an increase in uptake can be negated by concurrent increases in ecosystem C losses via respiratory processes. Continuous measurements of net ecosystem C exchange between the atmosphere and two experimental cheatgrass (Bromus tectorum L.) ecosystems in large dynamic flux chambers (EcoCELLs) showed net ecosystem C losses to the atmosphere in excess of 300 g C m^?2 over two growing cycles. Even a doubling of net ecosystem production (NEP) after N fertilization in the second growing season did not compensate for soil C losses incurred during the fallow period. Fertilization not only increased C uptake in biomass but also enhanced C losses through soil respiration from 287 to 469 g C m^?2, mainly through an increase in rhizosphere respiration. Fertilization decreased dissolved inorganic C losses through leaching of from 45 to 10 g C m^?2. Unfertilized cheatgrass added 215 g C m^?2 as root‐derived organic matter but the contribution of these inputs to long‐term C sequestration was limited as these deposits rapidly decomposed. Fertilization increased NEP but did not increase belowground C inputs most likely due to a concurrent increase in the production and decomposition of rhizodeposits. Decomposition of soil organic matter (SOM) was reduced by fertilizer additions. The results from our study show that, although annual grassland ecosystems can add considerable amounts of C to soils during the growing season, it is unlikely that they sequester large amounts of C because of high respiratory losses during dormancy periods. Although fertilization could increase NEP, fertilization might reduce soil C inputs as heterotrophic organisms favor root‐derived organic matter over native SOM.     

Evaluating contemporary pollen dispersal in two common grassland species Ranunculus bulbosus L. (Ranunculaceae) and Trifolium montanum L. (Fabaceae) using an experimental approach

Pollen flow is a key biological process that connects plant populations, preventing genetic impoverishment and inbreeding. Pollen‐mediated long‐distance dispersal (LDD) events are especially important for plant species in increasingly fragmented landscapes. Patterns of pollen dispersal were directly estimated and dispersal kernels modelled in an experimental population of Ranunculus bulbosus and Trifolium montanum to determine the potential for LDD. Eight and 11 microsatellite markers were used for R. bulbosus and T. montanum, respectively, to run a likelihood‐based paternity analysis on randomly chosen offspring (N_total = 180 per species) from five maternal plants. High rates of selfing were found in R. bulbosus (average 45.7%), while no selfing was observed in T. montanum. The majority (60%) of mating events occurred at very short distances: the median of the observed dispersal distances was 0.8 m in both species, and the average distances were 15.9 and 10.3 m in R. bulbosus and T. montanum, respectively. Modelling the pollen dispersal kernel with four different distribution functions (exponential‐power, geometric, 2Dt and Weibull) indicated that the best fit for both species was given by a Weibull function. Yet, the tail of the T. montanum pollen dispersal kernel was thinner than in R. bulbosus, suggesting that the probability for LDD is higher in the latter species. Even though the majority of pollen dispersal occurred across short distances, the detection of several mating events up to 362 m (R. bulbosus) and 324 m (T. montanum) suggests that pollen flow may be sufficient to ensure population connectivity in these herb species across fragmented grasslands in Swiss agricultural landscapes.