Ecological resistance, seed density and their interactions determine patterns of invasion in a California coastal grassland

Relatively little experimental evidence is available regarding how ecological resistance and propagule density interact in their effects on the establishment of invasive exotic species. We examined the independent and interactive effects of neighbour cover (biotic resistance), winter vs. spring water addition (abiotic resistance) and seed density on the invasion of the European perennial grass Holcus lanatus into a California coastal grassland dominated by exotic annual grasses. We found that decreased competition from resident exotic grasses had no effect. In contrast, increased late-season water availability eroded the abiotic resistance offered by naturally dry conditions, facilitating invasion. Finally, watering treatment and seed density interacted strongly in determining seedling survival: while seedling mortality was close to 100% in ambient and winter water addition plots, survivor numbers increased with seed density in spring-watered plots. Thus, decreased abiotic resistance can amplify the effect of increased propagule density on seedling establishment, thereby increasing the likelihood of invasion.     

Stochastic LTRE analysis of the effects of herbivory on the population dynamics of a perennial grassland herb

Herbivores can have strong deleterious effects on vital rates (growth, reproduction, and survival) and thus negatively impact the population dynamics of plant species. In practice, however, these effects might be strongly correlated, for example as a result of tradeoffs between vital rates. To get better insights into the effects of herbivory on the population dynamics of the long‐lived grassland plant Primula veris population projection matrices were constructed from demographic data collected between 1999 and 2008 (nine annual transitions). Data were collected in two large grassland populations, each of which was subjected to two treatments (grazing by cattle versus a mowing treatment), yielding a total of 36 matrices. We applied a lower‐level vital rate life table response experiment (LTRE) using the small noise approximation (SNA) of the stochastic population growth rate to disentangle the contributions of changes in mean vital rates, variability in vital rates, correlations between vital rates and vital rate elasticities to the difference in the stochastic growth rate. Stochastic growth rates (a= log λ_S) were significantly lower in grazed than in mown plots (a= 0.0185 and 0.1019, respectively). SNA LTRE analysis showed that contributions of mean vital rates by far made the largest contribution to the observed difference in a between grazed and control plots. In particular, changes in sexual reproduction rates made the largest contributions to lower the stochastic growth rate in grazed plots: both adult flowering probabilities and flower and seed production were importantly lower in grazed populations, but these negative effects were largely buffered by increased establishment and seedling survival rates. Among the stochastic terms of the SNA decomposition, contributions of covariance and correlations between vital rates had the largest impact, whereas contributions of elasticities were smaller. The strongest correlation driver was the association between adult survival and seedling establishment, suggesting that environmental conditions favouring adult survival also are beneficial for seedling establishment. Overall, our results show that herbivory had a strong negative effect on the long‐term population growth rate of P. veris that was primarily mediated by differences in fecundity (flower and seed production) and germination.     

Strain and vegetation effects on local limiting resources explain the outcomes of biotic interactions

Positive interactions are hypothesized to increase with stress (stress-gradient hypothesis, “SGH”), which is defined in terms of standing biomass at the community level. However, recent evidence suggests that facilitation may decrease or remain constant as stress increases. Several reasons for this discrepancy are possible: (i) the outcomes of biotic interactions depend on the component of the fitness considered; (ii) they are influenced by how vegetation affects local limiting resources; (iii) within a particular community, only species that are deviated from their physiological optima are likely to be facilitated. In a removal experiment, we quantified the deviations of species in subalpine grassland from their physiological optima, defined here as species-level “strain”, and examined whether strain and vegetation effects on local resources can explain the outcome of biotic interactions. The experiment was performed along a gradient of standing biomass driven by contrasting land use and resource availability, and used five grass species with contrasting traits and ecological optima. Strain for each species was estimated by comparing growth without vegetation (target species only submitted to local abiotic factors) to growth in optimal conditions (under controlled conditions in an experimental garden). The outcomes of biotic interactions, recorded in terms of survival and growth, could be predicted from the data about strain and vegetation effects on local limiting resources (light and water). Only highly strained species were affected by facilitation, which occurred when the surrounding vegetation alleviated the constraining factors. On the other hand, standing biomass was poorly related with the outcomes of biotic interactions. The “SGH” was only partially validated with growth data when strain and vegetation effects co-varied with standing biomass. As a consequence, strain (at species level) represents a mechanistic basis which could improve the prediction of the outcomes of biotic interactions along ecological gradients.     

Survival,growth, and photosynthesis of tree seedlings competing with herbaceous vegetation along a water-light-nitrogen gradient

In herbaceous dominated patches and ecosystems, tree establishment is influenced partly by the ability of woody seedlings to survive and grow in direct competition with herbaceous vegetation. We studied the importance of season long wet and dry spells on the competitive interactions between herbaceous vegetation and oak seedlings along a light and nitrogen gradient in an infertile secondary successional grassland in central North America. We conducted a field experiment in which seedlings of bur oak (Quercus macrocarpa) and northern pin oak (Q. ellipsoidalis) were exposed to two levels of light (full sun and 80% shade), three levels of nitrogen input (0, 5, 15 g m^–1 yr^–1), and three levels of water input (low, medium and high). In addition, seedlings were grown with and without the presence of surrounding herbaceous vegetation under both light and all three water levels. Seedling survival, growth, and rate of photosynthesis were significantly affected by competition with herbaceous vegetation and these effects varied along the multiple resource gradient. Overall, seedling survival of both species was significantly greater in wetter and shaded plots and when surrounding herbaceous vegetation was removed and was lower in nitrogen enriched plots. We found that soil water was significantly affected by varying inputs of water, light, and the presence or absence of herbaceous vegetation, and that seedling survival and rate of photosynthesis were highly correlated with available soil water. Our findings show that the impact of season long wet and dry spells on tree seedling success in grasslands can be affected by light and soil nitrogen availability.     

The contribution of understory light availability and biotic neighborhood to seedling survival in secondary versus old-growth temperate forest

Seedling survival plays an important role in the maintenance of species diversity and forest dynamics. Although substantial gains have been made in understanding the factors driving patterns of seedling survival in forests, few studies have considered the simultaneous contribution of understory light availability and the local biotic neighborhood to seedling survival in temperate forests at different successional stages. Here, we used generalized linear mixed models to assess the relative importance of understory light availability and biotic neighborhood variables on seedling survival in secondary and old-growth temperate forest in north eastern China at two levels (community and guild). At the community level, biotic neighborhood effects on seedling survival were more important than understory light availability in both forests. In both the old-growth and secondary forests, conspecific basal area had a negative effect on seedling survival, consistent with negative conspecific density dependence. At guild levels, the relative importance of light and biotic neighborhood on seedling survival showed considerable variation among guilds in both forests. Available understory light tended to have positive effects on seedling survival for shrub and light-demanding species in the old-growth forest, but negative effects on survival of shade-tolerant seedlings in the secondary forest. For tree species and shade-tolerant species, the best fit models included neighborhood variables, but that was not the case for shrubs, light-demanding, or mid shade-tolerant species. Overall, our results demonstrate that the relative importance of understory light availability and biotic factors on seedling survival vary with species life-history strategy and forest successional stage.     

Interactions between herbivory and resource availability on grazing tolerance of Leymus chinensis

Herbivory and resource interact to influence plant regrowth following grazing, but few detailed investigations on grazing tolerance at population levels are available. We conducted two pot experiments along a simulated grazing gradient (0%, 25%, 50% and 75% of shoot removal) at three water or nutrient levels to determine the interaction of resource and herbivory on Leymus chinensis, a perennial, dominant species in the eastern Eurasian steppes. Interactions between water availability and clipping intensity on the relative height growth rate (RHGR) and bud number were significant. Significant interactions between nutrient and clipping on RHGR, total biomass and specific leaf area (SLA) were also found. Total biomass and bud number, showing a unimodal curve along the clipping gradient in resource-rich environments, were highest at light clipping level, suggesting that this species has the plastic compensatory responses from under- to overcompensation. Interactions between herbivory and water or nutrient were opposite to each other. The “cooperative” interactions between water and herbivory magnified the difference in grazing tolerance of L. chinensis between high and low water treatments. The “antagonistic” interactions between nutrient and herbivory, on the other hand, were reflected in the lower tolerance to heavy clipping in the high nutrient than low nutrient treatments. Results partly support the limiting resource model (LRM). A modified and simplified graphic model of the LRM was proposed based on our results. The new LRM clearly demonstrated that “cooperative” interactions between varying water levels and clipping intensities aggravate the detrimental impacts of herbivores on plant growth and reproduction, whereas “antagonistic” interactions between nutrient and grazing alleviate the negative effects of herbivores. Biomass compensation and density compensation were identified as main mechanisms of herbivory tolerance in this clonal species.     

The role of biotic interactions in altering tree seedling responses to an extreme climatic event

Questions: We addressed two poorly understood aspects of plant response to climate change: the impact of extreme climatic events and the mediating role of biotic interactions, through a study of heatwave effects on tree seedling survival rates and ability of the tree canopy to alter seedling responses. Location: Mountain belt of the northern French Alps (Maurienne Valley). Methods: The survival rates of two seedling cohorts from four tree species (Abies alba, Acer pseudoplatanus, Fraxinus excelsior and Picea abies) were measured during both the 2003 European heatwave and an average summer (2004) in deciduous broadleaf mountain forests. Seedlings were transplanted into two soil moisture conditions, and in experimental gaps or under the tree canopy. Results: The heatwave strongly decreased tree seedling survival rates, while there was an important species‐specific mediating role of biotic interactions. In the wettest conditions, the tree canopy strongly increased survival of Abies, buffering the negative impact of the heatwave. In contrast, in the driest conditions, the tree canopy decreased survival of Picea and Acer, amplifying the negative impact of the heatwave. We found evidence of increasing soil water stress in the understorey of the driest community, but further studies including vapour pressure deficit measurements are needed to elucidate the driving mechanism of facilitation. Conclusions: The high species specificity of the mediating role of biotic interactions and its variation along stress gradients leads to questions on our ability to predict large‐scale responses of species to climate changes.     

Biotic Interactions Overrule Plant Responses to Climate,Depending on the Species' Biogeography

This study presents an experimental approach to assess the relative importance of climatic and biotic factors as determinants of species'' geographical distributions. We asked to what extent responses of grassland plant species to biotic interactions vary with climate, and to what degree this variation depends on the species'' biogeography. Using a gradient from oceanic to continental climate represented by nine common garden transplant sites in Germany, we experimentally tested whether congeneric grassland species of different geographic distribution (oceanic vs. continental plant range type) responded differently to combinations of climate, competition and mollusc herbivory. We found the relative importance of biotic interactions and climate to vary between the different components of plant performance. While survival and plant height increased with precipitation, temperature had no effect on plant performance. Additionally, species with continental plant range type increased their growth in more benign climatic conditions, while those with oceanic range type were largely unable to take a similar advantage of better climatic conditions. Competition generally caused strong reductions of aboveground biomass and growth. In contrast, herbivory had minor effects on survival and growth. Against expectation, these negative effects of competition and herbivory were not mitigated under more stressful continental climate conditions. In conclusion we suggest variation in relative importance of climate and biotic interactions on broader scales, mediated via species-specific sensitivities and factor-specific response patterns. Our results have important implications for species distribution models, as they emphasize the large-scale impact of biotic interactions on plant distribution patterns and the necessity to take plant range types into account.     

Growth of tree seedlings in a tropical dry forest in relation to soil moisture and leaf traits

"AimsThe growth of plant species in tropical dry forest (TDF) is expected to be largely governed by the availability of soil moisture. In this study we attempt to identify mechanisms by which seedlings of dry tropical trees cope with water stress by adjusting their leaf characteristics to water availability and micro environments, and address following questions: How are leaf traits and relative growth rate (RGR) of the dominant seedling species of TDF affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional traits with each other? Can leaf traits singly or in combination predict the growth rate of seedling species of TDF? The study was conducted in situ on four sites (viz., Hathinala, Gaighat, Harnakachar and Ranitali, listed in order of decreasing SMC) within the tropical dry deciduous forest in northern India. Methods Five leaf traits viz., specific leaf area (SLA), leaf dry matter content (LDMC), concentrations of leaf nitrogen (leaf N), phosphorus (leaf P) and chlorophyll (Chl) and two physiological processes, viz., stomatal conductance (Gs net) and photosynthetic rate (A net), and RGR, of four dominant tree seedling species of a TDF (viz., Buchanania lanzan, Diospyros melanoxylon, Shorea robusta and Terminalia tomentosa) on four sites were analysed for species, site and season effects over a 2-year period. Step-wise multiple regression was performed to predict RGR from mean values of SMC, leaf traits and physiological processes. Principal component analysis (PCA) was performed to observe the extent of intra- vs. inter-specific variability in the leaf traits and physiological rates.Important findings All the traits and physiological rates were interrelated and showed significant positive relationship with RGR except for the correlation of LDMC with RGR which was not significant. Further, relationships of SMC with all leaf traits, physiological rates and RGR were significant, except for that between SMC and SLA for B. lanzan and D. melanoxylon. The slope of seedling trait:SMC relationship, a measure of phenotypic plasticity in response to soil moisture gradient, varied among species. Among the four species, T. tomentosa was the most plastic and S. robusta the least. In conclusion, leaf traits and physiological processes were strongly related to soil water availability on the one hand and seedling growth on the other. Gs net is the most important variable which accounted for the greatest amount of variability (62%) in RGR, emphasizing the role of stomatal conductance in shaping growth patterns across spatial and temporal gradients of soil water availability. Gs net and SMC together explained 64% variability in RGR, indicating that other traits/factors, not studied by us are also important in modulating the growth of tropical tree seedlings.     

Direct and indirect interactions among plants explain counterintuitive positive drought effects on an eastern Mediterranean shrub species

Coexistence of woody and herbaceous plants may be governed by a complex set of direct and indirect interactions, whose relative importance have been rarely assessed. We experimentally studied woody species establishment in a mixed plant community by disentangling the potential role of such biotic interactions and the effect of environmental variations on them. Seedling establishment of the common eastern Mediterranean shrub species Sarcopoterium spinosum was investigated under different rainfall and light conditions, combined with the effect of the presence of adult shrubs and annual neighbors. We predicted that seedlings will be directly affected by competition with annuals with increasing water availability, while direct effects of adult shrubs will be positive via amelioration of water stress. Indirect effects were expected beneath shrub canopies due to reduced water stressed and light availability for both annuals and shrub seedlings, which may intensify competition between annuals and shrub seedlings. To test these predictions we performed field and garden experiments in which we combined manipulation of shrub and annual presence with manipulations of water availability and light conditions to simulate the effect of shrub canopy. In contrast to our prediction, shrub seedling establishment was not facilitated but inhibited by adult shrubs because of light limitation. As expected, annuals had direct negative effects on shrub seedlings under wet conditions, which shifted to neutral or positive effects under dry conditions. Thus, interactions among shrubs and annuals, and in particular the release from competition during drought years, leads to a counterintuitive positive effect of drought on shrub seedling establishment. Our findings point to the importance of experimentally studying multidimensional interactions for coexistence of different life forms and to the underestimated role of light for success in water‐limited ecosystems.     

Natural regeneration of selected timber species in the Republic of Congo

Natural regeneration of timber species is critical to the sustainable management of tropical forests. To understand what determines regeneration success of timber species in the Congo Basin, we evaluated whether seedling recruitment rates differed between forest logged 30 years previously and unlogged forest and determined the environmental factors that influence seedling density, growth and survival. We monitored the fate of 2186 seedlings of seven timber species within 462, 25‐m² plots located along 21 transects. We characterized seedling plots by light availability, soil nutrient availability and pH, and abundance of mammalian herbivores and then used linear and generalized linear mixed models to evaluate the variables that influenced seedling density, growth and survival. Light availability and canopy openness were 18% and 81% higher in logged than unlogged forest, and concentration of soil nutrients varied between sites. Seedling density was 32% higher in unlogged than logged forest. Taking all species together, seedling survival was positively correlated with calcium and negatively with magnesium and available phosphorus. Rates of seedling growth increased with available light. Taken separately, seedlings of the selected timber species responded differently to abiotic and biotic factors, demonstrating species‐specific regeneration requirements.     

Are variations of direct and indirect plant interactions along a climatic gradient dependent on species’ strategies? An experiment on tree seedlings

Investigating how interactions among plants depend on environmental conditions is key to understand and predict plant communities’ response to climate change. However, while many studies have shown how direct interactions change along climatic gradients, indirect interactions have received far less attention. In this study, we aim at contributing to a more complete understanding of how biotic interactions are modulated by climatic conditions. We investigated both direct and indirect effects of adult tree canopy and ground vegetation on seedling growth and survival in five tree species in the French Alps. To explore the effect of environmental conditions, the experiment was carried out at 10 sites along a climatic gradient closely related to temperature. While seedling growth was little affected by direct and indirect interactions, seedling survival showed significant patterns across multiple species. Ground vegetation had a strong direct competitive effect on seedling survival under warmer conditions. This effect decreased or shifted to facilitation at lower temperatures. While the confidence intervals were wider for the effect of adult canopy, it displayed the same pattern. The monitoring of micro‐environmental conditions revealed that competition by ground vegetation in warmer sites could be related to reduced water availability; and weak facilitation by adult canopy in colder sites to protection against frost. For a cold‐intolerant and shade‐tolerant species (Fagus sylvatica), adult canopy indirectly facilitated seedling survival by suppressing ground vegetation at high temperature sites. The other more cold tolerant species did not show this indirect effect (Pinus uncinata, Larix decidua and Abies alba). Our results support the widely observed pattern of stronger direct competition in more productive climates. However, for shade tolerant species, the effect of direct competition may be buffered by tree canopies reducing the competition of ground vegetation, resulting in an opposite trend for indirect interactions across the climatic gradient.     

Acacia karroo invasion of grassland: environmental and biotic effects influencing seedling emergence and establishment

Acacia karroo Hayne is the most important woody invader of grassland in South Africa, and can greatly reduce the productivity of grassland. A field experiment was conducted to test the hypotheses that emergence, growth and the 1st year's survival of Acacia karroo would be enhanced by (1) defoliation of the grass sward, (2) increased irradiance, (3) increased moisture availability and (4) its germination within cattle dung pats. The study was conducted on one site above and one below the natural altitudinal treeline of this species in grassland of the eastern Cape, South Africa. Not one seedling emerged from dung pats. Neither location nor the other treatments affected the density of emerging seedlings, although only 40.4 seedlings m^–2 emerged of the 200 seeds m^–2 planted. Shading dramatically increased the density of surviving seedlings. In the open, only 3 and 1.5 seedlings m^–2 remained respectively at the end of the growing season or the beginning of the next, compared to 23.3 and 19.5 seedlings m^–2 under shading for these respective times. This was attributed to the effect of shade on moisture availability in a season which received only 54% of average rainfall. Seedling survival until the end of the growing season was enhanced (30%) by shade at both sites, but also by supplemental water (24%) and defoliation of the sward (7%) at the site above the treeline. Across sites and treatments, seedling survival was related to moisture availability, with no or poor survival for < 500 mm rainfall, indicating this species can only establish in certain rainfall years. Seedling survival over winter was not influenced by treatment, but was greater for larger seedlings. Treatments affected seedling size, in particular seedlings growing under shade and within a dense grass canopy were etiolated. A. karroo seedlings are capable of establishing and surviving within a dense grass sward for at least a year, tolerant of low irradiance and of interference, which, because most seeds do not persist for much longer than a year, suggests this species forms predominantly a seedling bank. This has implications for the invasion of grassland by woody species.     

The confounding effects of logging on tree seedling growth and herbivory in Central Amazon

Light availability is an important modulator of seedling growth and plant–herbivore dynamics. Logging increases light levels in forests, potentially altering herbivore–plant interactions that drive seedling establishment. We conducted a transplant experiment to evaluate how logging and herbivory affect seedling growth and survival in three shade‐tolerant tree species, at paired canopy gap and understory sites in logged forest and an adjacent unlogged area in central Amazonia (Brazil). Seedlings were either left exposed to naturally occurring insect herbivores or protected from insects by a fine netting structure. We measured the herbivore damage and growth rate of seedlings after 18 mo. In general, logged areas received more light than unlogged sites. Growth and herbivory rates were positively influenced by light, and herbivory was also influenced positively by logging. In gaps, increased growth mitigated foliar damage. Logging resulted in a loss of foliar tissue due to increased herbivory. Herbivory rates were higher in the understory of logged sites than in that of unlogged understory sites, but growth was similar in these areas. Thus, the understory of logged areas provided the least favorable sites for shade‐tolerant tree regeneration, due to higher herbivory rates. The effect of logging on biotic interactions can extend beyond the gaps it creates into untouched understory sites. To our knowledge, this is the first time such a pattern has been observed, highlighting the importance of evaluating the impact of logging on biotic interactions.     

Resource allocation and storage relative to resprouting ability in wind disturbed coastal forest trees

Many plants persist by resprouting after disturbance. However, the benefits of resprouting (survival) may be traded off against height growth and reproduction. Resources (total non-structural carbohydrates—TNC) that could be allocated to growth or reproduction are stored or mobilised to support resprouting. TNC may either be stored by accumulation where availability exceeds the requirements for growth, or by reserve formation when storage is at the expense of growth. Thus, the mechanism of storage and resource allocation may differ between good (R+) and poor (R?) resprouters in response to nutrient availability and disturbance regime. R+ species typically reserve resources to ensure a rapid resprouting response to disturbance. We test whether R+ and R? species in coastal forest, under chronic wind disturbance, differ in growth rates, biomass allocation, leaf traits, water relations and storage of TNC. Seedlings from three confamilial pairs of R+ and R? tree species were subjected to nitrogen addition, water stress and clipping (simulating herbivory) treatments under greenhouse conditions. R? species had greater height growth rates, larger specific leaf area, lower root mass ratio and lower root TNC than R+ species. These differences between R+ and R? species were maintained irrespective of the levels of nitrogen, water and clipping treatments. R+ species did not increase their TNC concentration under nutrient and water stress, indicating that TNC is stored by reserve formation. R+ species appeared to trade-off growth against storage, while R? species did not. In R+ species, reserve formation is likely a bet-hedging strategy against occasional strong selection events in addition to chronic wind stress. By trading off height growth for better resprouting ability, good resprouters may be able to persist at more frequently disturbed sites (e.g., dune crests and windward slopes), while poor resprouters that have faster height growth can dominate less disturbed sites.     

The Effects of Established Trees on Woody Regeneration during Secondary Succession in Tropical Dry Forests

Understanding the mechanisms controlling secondary succession in tropical dry forests is important for the conservation and restoration of this highly threatened biome. Canopy‐forming trees in tropical forests strongly influence later stages of succession through their effect on woody plant regeneration. In dry forests, this may be complex given the seasonal interplay of water and light limitations. We reviewed observational and experimental studies to assess (1) the relative importance of positive and negative effects of established trees on regeneration; (2) the mechanisms underlying these effects; and (3) to test the ‘stress gradient hypothesis’ in successional tropical dry forests. The effects of established trees on seed dispersal, seed survival, and seed germination—either through direct changes to moisture and temperature regimes or mediated by seed dispersers and predators—are mainly positive. The balance between positive and negative effects on seedling establishment is more complex and depends on the season and leaf phenology of both trees and seedlings. Seedling survival is generally enhanced by established trees mitigating dry conditions. Established trees have counteracting effects on water and light availability that influence seedling growth. The probability of a positive effect of established trees on seedling survival decreases with increased rainfall, which supports the stress gradient hypothesis. Priorities for future research are experiments to test for facilitation and competition and their underlying mechanisms, long‐term studies evaluating how these effects change with ontogeny, and studies focusing on the species‐specificity of interactions.     

Buffering and plasticity in vital rates of oldfield rodents

1. Under the hypothesis of environmental buffering, populations are expected to minimize the variance of the most influential vital rates; however, this may not be a universal principle. Species with a life span <1 year may be less likely to exhibit buffering because of temporal or seasonal variability in vital rate sensitivities. Further, plasticity in vital rates may be adaptive for species in a variable environment with reliable cues. 2. We tested for environmental buffering and plasticity in vital rates using stage-structured matrix models from long-term data sets in four species of grassland rodents. We used periodic matrices to estimate stochastic elasticity for each vital rate and then tested for correlations with a standardized coefficient of variation for each rate. 3. We calculated stochastic elasticities for individual months to test for an association between increased reproduction and the influence of reproduction, relative to survival, on the population growth rate. 4. All species showed some evidence of buffering. The elasticity of vital rates of Peromyscus leucopus (Rafinesque, 1818), Sigmodon hispidus Say & Ord, 1825 and Microtus ochrogaster (Wagner, 1842) was negatively related to vital rate CV. Elasticity and vital rate CV were negatively related in Peromyscus maniculatus (Wagner, 1845), but the relationship was not statistically significant. Peromyscus leucopus and M. ochrogaster showed plasticity in vital rates; reproduction was higher following months where elasticity for reproduction exceeded that of survival. 5. Our results suggest that buffering is common in species with fast life histories; however, some populations that exhibit buffering are capable of responding to short-term variability in environmental conditions through reproductive plasticity.     

Effects of ramet clipping and nutrient availability on growth and biomass allocation of yellow nutsedge

A glasshouse experiment was conducted to examine how the interactions of nutrient availability and partial ramet clipping affect growth, reproduction and biomass allocation of Cyperus esculentus, an invasive sedge. The plants sprouting from tubers were grown at low and high nutrient levels, and were subject either to no clipping, one, two or three clippings, with each clipping cutting half of the existing ramets at soil level. Our results show that nutrient availability and clipping frequency tended to independently affect most of growth, reproduction and biomass allocation parameters of Cyperus esculentus examined in the present study. Increased supply of nutrients led to an increase in plant productivity and its associated traits. All of the traits, except for the number of ramets, displayed a decreasing pattern with increasing clipping frequency, indicating that Cyperus esculentus had undercompensatory responses to ramet clipping. It is likely that the patterns of plants response to clipping are species specific, and depend on morphological characters of species. Its susceptibility to ramet clipping can offer opportunities for controlling this invasive species through mechanical methods such as mowing. Clipping had little effects on biomass allocation; however, root weight fraction increased with increasing clipping frequency. While nutrient availability and clipping frequency had no influence on leaf carbon concentration at harvest, both of them increased leaf nitrogen concentration, and hence reduced leaf C/N ratio.     

Effects of below-ground insects, mycorrhizal fungi and soil fertility on the establishment of Vicia in grassland communities

 The effects of, and interactions between, insect root feeders, vesicular-arbuscular mycorrhizal fungi and soil fertility on the establishment, growth and reproduction of Vicia sativa and V. hirsuta (Fabaceae) were investigated in an early-successional grassland community. Seeds of both species were sown into plots where soil insecticide (Dursban 5G), soil fungicide (Rovral) and soil fertiliser (NPK) were applied in a factorial randomised block design. Fertiliser addition reduced growth, longevity and reproduction of both Vicia species, due to the commonly recorded increase in the competitive advantage of the non-nitrogen-fixing species when nitrogen is added to the plant community. However, in plots where fertiliser was not applied, a reduction in root feeders and mycorrhizal infection led to an increase in seedling establishment and fruit production of V. sativa, and to an increase in flower production for both Vicia species. The interaction between all three soil treatments explained much of the variation in growth and longevity of V. sativa. Plants grew larger and survived longer in plots where natural levels of mycorrhizal infection and root feeders were low compared with plots where all the treatments were applied. This suggests that, although soil nutrient availability was a strong determinant of the performance of these two leguminous species, at natural levels of soil fertility biotic factors acting in the soil, such as mycorrhizal fungi and soil-dwelling insects, were important in shaping the competitive interactions between the two Vicia species and the plant community. Our results indicate that non-additive interactions between ecological factors in the soil environment may strongly affect plant performance. Received: 18 July 1995 / Accepted: 14 August 1996     

Limits to tree species invasion in pampean grassland and forest plant communities

Factors limiting tree invasion in the Inland Pampas of Argentina were studied by monitoring the establishment of four alien tree species in remnant grassland and cultivated forest stands. We tested whether disturbances facilitated tree seedling recruitment and survival once seeds of invaders were made available by hand sowing. Seed addition to grassland failed to produce seedlings of two study species, Ligustrum lucidum and Ulmus pumila, but did result in abundant recruitment of Gleditsia triacanthos and Prosopis caldenia. While emergence was sparse in intact grassland, seedling densities were significantly increased by canopy and soil disturbances. Longer-term surveys showed that only Gleditsia became successfully established in disturbed grassland. These results support the hypothesis that interference from herbaceous vegetation may play a significant role in slowing down tree invasion, whereas disturbances create microsites that can be exploited by invasive woody plants. Seed sowing in a Ligustrum forest promoted the emergence of all four study species in understorey and treefall gap conditions. Litter removal had species-specific effects on emergence and early seedling growth, but had little impact on survivorship. Seedlings emerging under the closed forest canopy died within a few months. In the treefall gap, recruits of Gleditsia and Prosopis survived the first year, but did not survive in the longer term after natural gap closure. The forest community thus appeared less susceptible to colonization by alien trees than the grassland. We conclude that tree invasion in this system is strongly limited by the availability of recruitment microsites and biotic interactions, as well as by dispersal from existing propagule sources.