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Contrasting phenotypes of foundation species are known to differentially affect understorey plants. However, there is little knowledge on both the mechanisms of competition (resource competition versus interference) of stress‐tolerant phenotypes and the importance of indirect interactions. In an oromediterranean community from Mount Lebanon we assessed the effects on understorey forbs of two contrasting phenotypes, a tight competitive from stressful habitat and a loose facilitative from more benign habitat. In a dry south and a wet north exposure we assessed short‐term resource effects removing shrub canopy and long‐term soil effects (including litter interference) with the comparison of forb performances in adjacent naturally open areas vs no shrub. Indirect effects were quantified through the removal of grasses. Abundance, richness and biomass of forbs were measured in all treatments after one year of experiment, together with litter depth and soil moisture. We found strong direct negative soil effects of the tight phenotype on all forb performances and in particular in south exposure. These effects were due to litter interference on water availability, but not to resource competition. They were likely explained by the high hydrophobicity of organic matter accumulating in the stressful habitat of the tight phenotype. We also found an indirect competition of the loose phenotype for forb richness, due to its direct positive soil effect on competitive grasses, and in particular in south exposure. Our results improve our knowledge on the importance of litter interference in dry nutrient‐poor habitats and the role of indirect interactions in phenotypic effects on understorey species.

The phenology of seed ripening and release are important for dispersal, reproductive success and survival of plants. Most phenological studies, however, consider early phenological phases. Here, we examined the ecological and evolutionary basis of ripening and seed release phenology. We monitored single flower phenology for 104 plant species from 30 families and three life forms from central Europe. Further, we undertook an associate monitoring study along an elevational gradient over two years. We calculated temperature demands (as growing degree days) for ripening and seed release and examined them with respect to the species’ seed mass, life form, dispersal mode and phylogeny. We found a strong correlation between species’ seed mass and temperature demands for ripening. For both variables seed mass and temperature demands for seed ripening, we found a strong effect of the species phylogeny. These phylogenetic signals indicate that the evolutionary history of the species’ lineage affects its seed mass and the temperature demands for seed ripening. Among the studied life forms, shrub species showed the most efficient ripening process. Anemochorous species showed lower relative humidity during seed release than epizoochorous species. For anemochorous species, the synchronisation of release timing with periods that show favourable environmental conditions for wind dispersal could be interpreted as a phenological adaptation to increase dispersal distances. According to the monitoring along the elevational gradient, individuals from higher altitudes showed lower temperature demands for ripening than individuals from lower altitudes. This might tentatively indicate physiological adaptations to lower temperature demands for locations with a shorter growing season. Our study provides basic insights into the ecological, environmental and evolutionary constraints that shape the ripening and seed release phenology of plants. We introduce data that can be used to advance existing models of ripening phenology, seed release and plant spread.

A necessary condition for a snowshoe hare population to cycle is reduced reproduction after the population declines. But the cause of a cyclic snowshoe hare population’s reduced reproduction during the low phase of the cycle, when predator density collapses, is not completely understood. We propose that moderate-severe browsing by snowshoe hares upon preferred winter-foods could increase the toxicity of some of the hare’s best winter-foods during the following hare low, with the result being a decline in hare nutrition that could reduce hare reproduction. We used a combination of modeling and experiments to explore this hypothesis. Using the shrub birch Betula glandulosa as the plant of interest, the model predicted that browsing by hares during a hare cycle peak, by increasing the toxicity B. glandulosa twigs during the following hare low, could cause a hare population to cycle. The model’s assumptions were verified with assays of dammarane triterpenes in segments of B. glandulosa twigs and captive hare feeding experiments conducted in Alaska during February and March 1986. The model’s predictions were tested with estimates of hare density and measurements of B. glandulosa twig growth made at Kluane, Yukon from 1988–2008. The empirical tests supported the model’s predictions. Thus, we have concluded that a browsing-caused increase in twig toxicity that occurs during the hare cycle’s low phase could reduce hare reproduction during the low phase of the hare cycle.

Direct and indirect interactions among plants contribute to shape community composition through above- and belowground processes. However, we have not disentangled yet the direct and indirect soil and canopy effects of dominants on understorey species. We addressed this issue in a semi-arid system from southeast Spain dominated by the legume shrub Retama sphaerocarpa. During a year with an exceptionally dry spring, we removed the shrub canopy to quantify aboveground effects and compared removed-canopy plots to open plots between shrubs to quantify soil effects, both with and without watering. We added a grass removal treatment in order to separate direct from indirect shrub effects and quantified biomass, abundance, richness and composition of the forb functional group. With watering, changes in forb biomass were primarily driven by indirect shrub effects, with contrasting negative soil and positive aboveground indirect effects; changes in forb abundance and composition were more influenced by direct shrub soil effects with contrasting species composition between open and Retama patches. As community composition was different between open and Retama patches the indirect effects of Retama on forb species did not concern forbs from the open community but forbs from Retama patches. Indirect effects are, thus, important at the functional group level rather than at the species level. Without watering, there were no significant interactions. Changes in species richness between treatments were weak and seldom significant. We conclude that shrub effects on understorey forbs are primarily due to their influence on soil properties, directly affecting forb species composition but indirectly affecting the biomass of the forbs of the Retama patches, and only with sufficient water.

Facilitation by nurse plants plays an important role in determining community composition in severe environments. Although the unidirectional effect of nurses on beneficiary species has received considerable research interest, nurse-mediated interactions among beneficiary species (so-called indirect interactions) are less known. Consequently, community composition in nurse plant systems is generally considered as a simple consequence of the facilitative effect of the nurse even though beneficiary species may significantly contribute to community assembly and modulate the direct nurse effects on the community. In an observational study we assessed nurse effects and nurse-mediated beneficiary interactions in two contrasting nurse plant systems in dry environments using a newly developed framework. We quantified plant–plant interaction intensity using the relative interaction index (RII) at the community and species level for three Retama sphaerocarpa shrub size-classes in a semiarid shrubland and four Arenaria tetraquetra cushion plant communities differing in aspect and elevation in dry alpine gravel habitats. The observed RII was split into nurse and beneficiary effects, and related to individual mass, species frequency and abundance using generalized linear mixed models. Results showed predominantly positive nurse effects and negative beneficiary interactions. The effect size of nurse plants, however, was significantly higher than the effect size of beneficiary species in both systems. Individual plant mass and abundance of species was dependent on the combined effects of nurse and beneficiary species whereas species occurrence was related to nurse effects only. Despite evident differences, the semiarid and alpine nurse plant systems showed strong functional parallelisms. We found interdependence between the effects of nurse and beneficiary species on beneficiary plant assemblages emphasizing their combined role on community assembly in both systems. Our results highlight the need to consider indirect interactions to understand fully plant community dynamics.

Plant and animal survey detection rates are important for ecological surveys, environmental impact assessment, invasive species monitoring, and modeling species distributions. Species can be difficult to detect when rare but, in general, how detection probabilities vary with abundance is unknown. We developed a new detectability model based on the time to detection of the first individual of a species. Based on this model, the predicted detection rate is proportional to a power function of abundance with a scaling exponent between zero and one that depends on clustering of individuals. We estimated the model parameters with data from three independent datasets: searches for chenopod shrub species and coins, experimental searches for planted seedlings, and frog surveys at multiple sites in sub-tropical forests of eastern Australia. Analyses based on the detection time and detection probability suggest that detection rate increases with abundance as predicted. The model provides a way to scale detection rates to cases of low abundance when direct estimation of detection rates is often impractical.

Human driven changes in land-use have increased the need to understand how landscape structure affects species distribution. We studied how forest edges affected the distribution of birds in grasslands recently encroached by forest patches. We investigated how species’ biological traits influenced their response to vegetation change near forest edges. We censured birds along 300-m line transects run into the open habitat perpendicularly to forest edges. We recorded habitat variables and landscape context along each transect and characterized edges and forest patches. We recorded 33 bird species in 153 transects for a total of 654 individuals. We analyzed species response to edges with generalized linear mixed models. Habitat preference was prevalent to explain species response to forest edges. The abundance of open-habitat birds such as skylark Alauda arvensis decreased significantly in the vicinity of edges. This negative response extended within 150 m from the edge. The effect was disproportionately higher in open-habitat species with high conservation concern. The abundance of species feeding or/and breeding in both forest and open habitat, such as woodlarks Lullula arborea, sharply increased near edges (positive edge response). Abundance of shrub and non-shrub dependent species increased with distance to edge. The two species groups did no differ in abundance/distance to edge relationship. Intensity of species response to forest edges varied among transects in relation to transect vegetation characteristics. Edge length or aspect, diet and nest height had no direct effect. We discuss the possible role of variation in resources and nest predation risk to explain observed patterns.

Desertification threatens 70% of all dry lands worldwide by diminishing the provision of economic and ecosystem services. However, since long-term vegetation dynamics of semiarid ecosystems are difficult to study, the opportunities to evaluate desertification and degradation properly are limited. In this study, we tailored, calibrated and tested a spatially-explicit simulation model (DINVEG) to describe the long-term dynamics of dominant grass and shrub species in the semiarid Patagonian steppe. We used inverse techniques to identify parameterizations that yield model outputs in agreement with detailed field data, and we performed sensitivity analyses to reveal the main drivers of long-term vegetation dynamics. Whereas many parameterizations (10–45%) matched single field observations (e.g. grass and shrub cover, species-specific density, aboveground net primary production [ANPP]), only a few parameterizations (0.05%) yielded simultaneous match of all field observations. Sensitivity analysis pointed to demographic constraints for shrubs and grasses in the emergence and recruitment phase, respectively, which contributed to balanced shrub-grass abundances in the long run. Vegetation dynamics of simulations that matched all field observations were characterized by a stochastic equilibrium. The soil water content in the top layer (0–10 cm) during the emergence period was the strongest predictor of shrub densities and population growth rates and of growth rates of grasses. Grasses controlled the shrub demography because of the resource overlap of grasses with juvenile shrubs (i.e. water content in the top layer). In agreement with field observations, ecosystem function buffered the strong variability in precipitation (a simulated CV in ANPP of 16% vs CV in precipitation of 33%). Our results show that seedling emergence and recruitment are critical processes for long-term vegetation dynamics in this steppe. The methods presented here could be widely applied when data for direct parameterization of individual-based models are lacking, but data corresponding to model outputs are available. Our modeling methodology can reduce the need for long-term data sets when answering questions regarding community dynamics.

Changes in climate and in browsing pressure are expected to alter the abundance of tundra shrubs thereby influencing the composition and species richness of plant communities. We investigated the associations between browsing, tundra shrub canopies and their understory vegetation by utilizing a long-term (10–13 seasons) experiment controlling reindeer and ptarmigan herbivory in the subarctic forest tundra ecotone in northwestern Fennoscandia. In this area, there has also been a consistent increase in the yearly thermal sum and precipitation during the study period. The cover of shrubs increased 2.8–7.8 fold in exclosures and these contrasted with browsed control areas creating a sharp gradient of canopy cover of tundra shrubs across a variety of vegetation types. Browsing exclusions caused significant shifts in more productive vegetation types, whereas little or no shift occurred in low-productive tundra communities. The increased deciduous shrub cover was associated with significant losses of understory plant species and shifts in functional composition, the latter being clearest in the most productive plant community types. The total cover of understory vegetation decreased along with increasing shrub cover, while the cover of litter showed the opposite response. The cover of cryptogams decreased along with increasing shrub cover, while the cover of forbs was favoured by a shrub cover. Increasing shrub cover decreased species richness of understory vegetation, which was mainly due to the decrease in the cryptogam species. The effects were consistent across different types of forest tundra vegetation indicating that shrub increase may have broad impacts on arctic vegetation diversity. Deciduous shrub cover is strongly regulated by reindeer browsing pressure and altered browsing pressure may result in a profound shrub expansion over the next one or two decades. Results suggest that the impact of an increase in shrubs on tundra plant richness is strong and browsing pressure effectively counteracts the effects of climate warming-driven shrub expansion and hence maintains species richness.

Treelines have drawn persistent research interest as they can respond markedly to climate. However, the mechanisms that determine tree seedling recruitment and the response of the forest-tundra ecotone to environmental changes remain poorly understood. We hypothesise that treeline tree seedling performance depends on the interplay between climatic and soil nutritional changes and facilitative and competitive interactions between trees and shrubs. We conducted a seedling transplantation experiment with Betula pubescens at a subarctic treeline, in northern Sweden, which followed a full factorial design with four treatment factors relating to environmental regimes of stress and resource availability: site (forest vs treeline); temperature (+/- passive warming); shrub presence (+/-Vaccinium myrtillus removal); and nutrient availability (+/- NPK addition). During three growing seasons we assessed the establishment and performance of Betula. The experimental manipulations caused highly significant effects on seedling performance. Although Vaccinium enhanced seedling survival and reduced the effects of excessive solar radiation and insect herbivory, the seedlings growing with the shrub had a poorer performance by the end of the experimental period. Also, seedlings in the forest had a poorer performance than those at the treeline. Betula seedlings showed a very pronounced and positive response to passive warming and to nutrient addition, but such effects were more evident at the treeline site and often interacted with the presence of Vaccinium. This experiment shows that shrub-tree interactions are important drivers of subarctic treeline dynamics and that they vary with time and space. Facilitation, competition, herbivory and environmental changes at the tree seedling stage act as important filters in structuring the forest-tundra ecotone. We demonstrate that changes in this ecotone cannot be simply predicted from changing temperature patterns alone, and that complex interactions need to be considered, not only between shrubs and trees, but also with herbivores and between warming and soil nutrient availability. © 2012 The Authors.

Positive and negative plant-plant interactions are major processes shaping plant communities. They are affected by environmental conditions and evolutionary relationships among the interacting plants. However, the generality of these factors as drivers of pairwise plant interactions and their combined effects remain virtually unknown. We conducted an observational study to assess how environmental conditions (altitude, temperature, irradiance and rainfall), the dispersal mechanism of beneficiary species and evolutionary relationships affected the co-occurrence of pairwise interactions in 11 Stipa tenacissima steppes located along an environmental gradient in Spain. We studied 197 pairwise plant-plant interactions involving the two major nurse plants (the resprouting shrub Quercus coccifera and the tussock grass S. tenacissima) found in these communities. The relative importance of the studied factors varied with the nurse species considered. None of the factors studied were good predictors of the co-ocurrence between S. tenacissima and its neighbours. However, both the dispersal mechanism of the beneficiary species and the phylogenetic distance between interacting species were crucial factors affecting the co-occurrence between Q. coccifera and its neighbours, while climatic conditions (irradiance) played a secondary role. Values of phylogenetic distance between 207-272.8 Myr led to competition, while values outside this range or fleshy-fruitness in the beneficiary species led to positive interactions. The low importance of environmental conditions as a general driver of pairwise interactions was caused by the species-specific response to changes in either rainfall or radiation. This result suggests that factors other than climatic conditions must be included in theoretical models aimed to generally predict the outcome of plant-plant interactions. Our study helps to improve current theory on plant-plant interactions and to understand how these interactions can respond to expected modifications in species composition and climate associated to ongoing global environmental change.

Temperate forests are one of the most important ecosystems in the world, and thus disentangling the factors that drive diversity within these ecosystems is of major concern. However, due to the complex interactions among forests layers, topography and soil factors, discovering the drivers of diversity is often complicated. In this study, we tested three a priori hypotheses about the effect of the dominant competitor (Pinus koraiensis) on the different forest layers in a 25 ha full mapped plot of temperate forest in the Changbai Mountain of northeastern China. Structural equation modelling (SEM) was used to study the direct and indirect interactions between four vertical forest layers (dominant competitor, canopy composition, sub-canopy diversity and shrub diversity), topographic factors, edaphic factors to discover sub-canopy and shrub diversity drivers. Our results suggest that the dominant competitor (Pinus koraiensis) is a key factor explaining canopy variation, and sub-canopy diversity and shrub diversity, and that this competitor can act directly (through shading) and indirectly (through the modification of the soil). Topographic heterogeneity also had significant effects on the soil variation and the diversity of the sub-canopy and shrub layers. Finally our results indicate that the influence of canopy composition on the diversity of the rest of forest layers is indirect and positive, suggesting that the dominant competitor is the main factor limiting diversity. In conclusion, we have found strong evidence that the dominant species of the canopy can influence, both directly and indirectly, the diversity of the different vertical forest layers. Patterns of diversity in forests are driven by a multiplicity of factors that are inherently related.

The two major disturbance types of boreal black spruce forest in north–central Quebec, Canada – natural disturbance by wildfire and anthropogenic disturbance by harvest – may affect processes of recovery differently and leave distinct post‐disturbance soil and vegetation spatial patterns. We tested whether 1) spatial patterns of physico‐chemical soil organic layer properties, black spruce diameter and density, and understory ericaceous shrub cover, differ between these two principal disturbance types; 2) operations associated with forest harvest result in distinct, regular spatial patterns of these same variables related to presence of machine trails; and 3) ericaceous shrub presence is a potential factor contributing to the legacy of spatial patterns after harvest. We explored these patterns on black spruce‐feathermoss forest stands, including fire‐origin stands (18 and 98 years) and stands originating from harvest (16 and 62 years) in central Quebec, Canada. We used two spatial analysis methods, spectral analysis and principal component analysis in the frequency domain, to characterize and relate spatial patterns of these soil and vegetation variables, measured along 50‐m transects on each site. Spatial patterns of distribution of soil and vegetation variables were different on the burned and the harvested forest sites. Wildfire gave rise to spatial patterns in soil and vegetation variables at multiple scales, reflecting the complexity generated by variable burn intensity. Patterns following forest harvest were mainly related to the regular structure defined by trails created by logging operations. In contrast to burned sites, ericaceous shrub patterns on harvested sites were strongly associated with spatial arrangements of spruce diameter and density, promoting absence of canopy closure and persistence of trails. Moreover, different spatial signatures did not converge in the long‐term (62–98 years) between the two disturbance types. The divergence in spatial structure between natural and anthropogenic disturbances has implications for ecosystem structure and function in the longer term.

Understanding changes in community composition caused by invasive species is critical for predicting effects on ecosystem function, particularly when the invasive threatens a foundation species. Here we focus on dynamics of forest structure, composition and microclimate, and how these interact in southern Appalachian riparian forests following invasion by hemlock woolly adelgid, HWA, Adelges tsugae. We measured and quantified changes in microclimate; canopy mortality; canopy and shrub growth; understory species composition; and the cover and diversity in riparian forests dominated by eastern hemlock Tsuga canadensis over a period of seven years. Treatments manipulated hemlock mortality either through invasion (HWA infested stands) or girdling (GDL) hemlock trees. Mortality was rapid, with 50% hemlock tree mortality occurring after six years of invasion, in contrast to more than 50% mortality in two years following girdling. Although 50% of hemlock trees were still alive five years after infestation, leaf area lost was similar to that of girdled trees. As such, overall responses over time (changes in light transmittance, growth, soil moisture) were identical to girdled stands with 100% mortality. Our results showed different growth responses of the canopy species, shrubs and ground layer, with the latter being substantially influenced by presence of the evergreen shrub, rhododendron Rhododendron maximum. Although ground layer richness in the infested and girdled stands increased by threefold, they did not approach levels recorded in hardwood forests without rhododendron. Increased growth of co-occurring canopy trees occurred in the first few years following hemlock decline, with similar responses in both treatments. In contrast, growth of rhododendron continued to increase over time. By the end of the study it had a 2.6-fold higher growth rate than expected, likely taking advantage of increased light available during leaf-off periods of the deciduous species. Increased growth and dominance of rhododendron may be a major determinant of future responses in southern Appalachian ecosystems; however, our results suggest hemlock will be replaced by a mix of Acer, Betula, Fagus and Quercus canopy genera where establishment is not limited by rhododendron.

Defoliation through herbivory is well known to affect target plants and their associated belowground properties, but the response of plants and their soil environment to defoliation of their neighbours is less well understood. We performed a controlled shade-house experiment involving three plant species that colonize New Zealand floodplains during primary succession, i.e. a palatable N2-fixing shrub (Carmichaelia odorata), a palatable deciduous small tree (Fuchsia excorticata) and a less palatable evergreen tree (Weinmannia racemosa). All species were grown in large pots for 40 months both singly and in two species pairs, and either one or both of the species grown in pairs were clipped to simulate herbivory. Responses of growth and foliar nutrient status to clipping varied strongly among species, with Carmichaelia having the largest response and Fuchsia having the smallest. Carmichaelia also enhanced soil microbial biomass and activity, and foliar N concentrations of Weinmannia. However, this did not translate to a net positive effect; instead Carmichaelia competitively reduced growth and foliar P concentrations of both other species. Most effects of Carmichaelia on the soil microflora, and growth and nutrient status of its neighbours, disappeared when Carmichaelia was clipped. Further, the effect of clipping Carmichaelia had a stronger impact on growth, soil activity and nutrient status of the other two species than did the clipping of those species. These results contradict expectations that N2-fixing plants should promote growth of other species in pioneer communities or that defoliation of N2-fixers exacerbate positive effects; in our study, defoliation of Carmichaelia merely mitigated the negative effects that it had on other species. They also suggest that interplay of competition and differential herbivory among coexisting plants has important implications for soil microflora and processes, relative nutrient acquisition and stoichiometry of coexisting plant species, and potentially plant community development.

Th e study of plant – plant interactions along grazing and abiotic stress gradients is a major research topic in plant ecology, but the joint eff ects of both stressors on the outcome of plant – plant interactions remains poorly understood. We used two diff erent factorial experiments conducted in a semi-arid Mediterranean steppe to assess: 1) the role of the perennial grass Stipa tenacissima , a low-palatability species, providing protection from rabbit herbivory to the shrub Retama sphaerocarpa (experiment 1), and 2) the eff ects of environmental amelioration provided by Stipa on the recovery of Retama after rabbit damage under two contrasted levels of water availability (experiment 2). In the experiment 1, water stress worked as an indirect modulator of herbivore protection by Stipa. Th is species protected Retama seedlings from rabbit herbivory during the wetter conditions of spring and winter, but this eff ect dissapeared when rabbit pressure on Retama increased during summer drought due to the decrease in alternative food resources. In the experiment 2, Stipa exerted a negative eff ect on the survival of Retama seedlings during the three years of the experiment, regardless of inter-annual diff erences in rainfall or the watering level applied. Th is negative eff ect was mainly due to excessive shading. However, Stipa increased Retama recovery after initial rabbit impact, overriding in part this negative shade eff ect. Conversely, Stipa impact on the Fv/Fm of Retama seedlings depended on the intra-annual water dynamics and its experimental manipulation, overall contradicting predictions from the stress – gradient hypothesis. Th e complex interactions found between herbivory, microclimatic amelioration from Stipa , and water availability as drivers of Retama performance illustrate the importance of considering the temporal dynamics of both biotic and abiotic stressors to fully understand the outcome of plant – plant interactions.

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.

Plant populations are subjected to changes in their natural environment as a result of the incidence of simultaneous global change drivers. Despite the fact that these changes can largely affect early fitness components, information on the effects of simultaneous drivers of global change on offspring traits and performance is particularly scant. We analyzed the combined effect of three global change drivers of critical importance in Mediterranean ecosystems (habitat fragmentation, reductions in habitat quality and water availability) on germination and seedling performance of the gypsophile shrub Centaurea hyssopifolia. Seedlings from 39 mother plants from eight different environments (resulting from the combination of the three global change drivers) were sown and grown in a common garden. First, germination percentage, seedling size and seedling survival were monitored. Secondly, seedling performance and ecophysiological traits were measured under well and low-watered conditions. Fragmentation showed the largest negative effect on germination and offspring performance. Seedlings of mothers from small fragments germinated more slowly, showed lower survival, died faster, and showed lower photosynthetic rates under well-watered conditions compared to seedlings of mother plants from large populations. Seedlings of different maternal origins did not differ in their response to water stress or in their ability to survive to drought. Ninety-five percent of the seedlings survived until soil water content was as low as 3%. Our study shows that global change can have not only immediate impacts on plant populations but also transgenerational effects, and highlights the importance of studies involving multiple drivers and a more integral understanding of global change.

Following disturbance events vegetation can either be resilient and return to its original state, or there can be shifts in vegetation composition and abundance patterns that may indicate alternative equilibiria. We conducted a long-term field experiment that simulated impact by aerially transported volcanic ejecta (tephra) in a Sphagnum-dominated plant community in order to test the effects of this type of large-scale disturbance. Sixty plots of 1.4 x 1.4 m were established at Sarobetsu mire in northern Hokkaido, Japan and subjected to seven treatments (including the control) with natural tephra or ground glass shards differing in grain size, layer thickness and season of application. Water chemistry and vegetation were surveyed before tephra application and during five and eight years after the perturbation, respectively. Leaching of ions from fine-grained glass shards caused a sustained increase of soil water pH and electric conductivity. Under coarser materials water quality differed little from the control, but a short-lived peak of potassium suggested that mechanisms like nutrient release from decomposing plant material may influence water chemistry after volcanic disturbance. The perturbation initially reduced the cover of the dominant functional group (Sphagnum mosses) in all treatments; vascular plants were less affected. All species were able to recover by growing through the tephra, and open tephra surfaces were colonized by ubiquitous cryptogams, but not by spermatophytes. In contrast to the overall resilient behaviour of the vegetation, in some plots that received natural tephra an alternative state with high cover of the dwarf shrub Myrica gale developed. The patterns indicated that physical and chemical properties of the tephra determined the initial effects on plants, but that stochastic processes contributed to subsequent succession. These are likely to have effects on ecosystem functioning, e.g. hydrological processes and carbon sequestration.

savannah, we established two plots (respectively 550 and 500 m 2 ) within which we mapped all surviving and dead shrubs as well as all individuals of shrub species that recruited in the following year. We used techniques of point-pattern analysis to test specific null hypotheses about spatial associations in the distribution, mortality, and establishment of shrubs. Results support the notions that fire mediates interactions between these two species. Fire-caused death of E. buniifolium tended to occur selectively in the vicinities of Baccharis individuals, and recruitment of B. dracunculifolia tended to be concentrated in the places of dead shrubs. These responses, however, were contingent on local abundances of shrubs which depend in part from the recent fire history. Anthropogenic perturbation of the natural fire regime would have therefore distorted the role of fire mediated interactions as drivers of the dynamics of the vegetation of this temperate savannah.

Theoretically, induced defenses should be prevalent within low resource environments like the forest understory where constitutive defenses would be costly. Also, the induced response should be stronger when the herbivore is a generalist rather than a specialist, which often have mechanisms to avoid or overcome plant defenses. These ideas have been previously tested for herbaceous species, and we examined these predictions in Lindera benzoin (spicebush), a common woody shrub of the eastern deciduous forest. Lindera benzoin plants in contrasting light environments served as control plants or were subjected to one of four treatments: application of jasmonic acid, clipping, herbivory by the specialist Epimecis hortaria (tulip tree beauty) and herbivory by the generalist Spodoptera exigua (beet armyworm). Following treatment, we assessed induced responses by measuring leaf chemistry (C/N ratio, protein content, and peroxidase activity), and by using insect bioassays with E. hortaria larvae. We found no difference in peroxidase activity between light environments in controls, plants treated with clipping or jasmonic acid. In plants subject to insect herbivory, peroxidase activity was greater in shade plants than in sun plants. The magnitude of this increase in the shade varied between the herbivores, with a 32 fold increase in plants exposed to the generalist S. exigua and a 9 fold increase in plants exposed to the specialist E. hortaria. Leaves from shade plants had more protein and lower C/N ratios than leaves from sun plants, regardless of induction treatment. In control plants, E. hortaria larvae consumed more leaf biomass and achieved greater final weights in the sun than in the shade, but these differences disappeared with induction treatments were applied. These results are among the first to show rapid induction in a woody plant, and different levels of induction with light environments and with specialist versus generalist herbivores.

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.

Plant facilitation studies commonly test the nurse-plant hypothesis wherein an adult shrub species enhances the establishment of associated herbaceous species under its canopy. Using field and glasshouse experiments, this hypothesis is extended by testing the following four predictions: (1) nurse-plant effects can occur between species with similar life-forms and phenologies (2) positive effects are species specific, (3) the outcome of interactions is life-stage dependent, and (4) facilitative interactions among annuals are primarily commensal. In the Negev Desert in Israel, the response of an annual plant community to removal of relatively larger annuals, Erodium laciniatum, Erucaria pinnata and Trifolium tomentosum, was tested in the field and in the glasshouse. Removal of these dominants was applied early in the growing season, immediately after germination but before establishment of seedlings, and again mid-season following establishment and growth to adults. In both the field and glasshouse, the presence of E. laciniatum increased establishment and survival to reproduction of neighbouring plants. These positive effects were life-stage dependent with more positive effects occurring early in the season, and there was no cost of facilitation to E. laciniatum. This positive effect was species specific in that neither E. pinnata nor T. tomentosum had an effect on the plant community. There was also a cost of association with the nurse E. laciniatum in that biomass of neighbours was reduced. These experiments demonstrate that the positive effects typically detected at larger scales between species of different life-forms are also occurring at finer spatial scales amongst annuals. This study clearly supports the predictions made in the facilitation literature that effects are species-specific and highly life-stage dependent.

The fitness benefits of plant structural adaptations that increase the effectiveness of fungivores against leaf pathogenic fungi are poorly understood. In a 12-month field experiment, we investigated the effect of domatia on mite density, the role of these mites in limiting leaf fungi, and the associated effects on plant fitness in the endemic New Zealand shrub, Coprosma lucida. The presence of domatia on mite density was controlled using combinations of domatia blocking, sham blocking, mite addition and mite control using miticide. Limiting access to domatia reduced mite density and increased the proportion of leaves without mites. Mite families represented were predominantly fungivorous/detritivorous (97.2%), and predaceous (2.6%); herbivorous mites were absent. Mites significantly reduced fungal hyphae, fungal spores and pollen, but the effect was surface(upper/lower) and density-dependent with the greatest reduction in fungi occurring over low mite densities. Fungal hyphae reduced leaf longevity, but were associated with increased production of new leaves. Hyphae density on old leaves was negatively correlated with the number of domatia produced on new leaves. New leaves in the mite reduction treatment had slightly reduced levels of carbon but not nitrogen. High levels of fungal infection on the lower surface increased the number of fruit fascicles per shoot, however on the upper surface where fungi were reduced by mites, hyphae density was negatively related to reproduction. The data support a limited interpretation of a fitness benefit for plants with domatia. While domatia increased mite density, control of fungi by mites occurred at lower average densities than supported by plants without functioning domatia. We suggest the primary function of leaf domatia in this mutualism is to increase the probability of a leaf-level beneficial mite presence rather than to maximise mite density. Many mites are not necessarily better than few mites, but some mites are better than none. Plant-directed defence mutualisms involving invertebrates require facilitation of the presence of beneficial

Spatial distribution of palatable and unpalatable plants can influence the foraging behaviour of herbivores, thereby changing plant-damage probabilities. Moreover, the immediate proximity to certain plants can benefit other plants that grow below them, where toxicity or spines act as a physical barrier or concealment against herbivores. This paper presents the results of a multi-scale experiment performed to test the effect of shrubs as protectors of tree saplings against herbivores and the mechanism involved in Mediterranean ecosystems. We performed a factorial design in two mountain ranges, similar in physiognomy and vegetation, planting saplings of a palatable tree, the maple (Acer opalus subsp. granatense), and an unpalatable tree, the black pine (Pinus nigra), under three different types of shrubs. We considered four experimental microhabitats: highly palatable shrub (Amelanchier ovalis), palatable but spiny shrub (Crataegus monogyna or Prunus ramburii), unpalatable spiny shrub (Berberis vulgaris subsp. australis) and control (gaps of bare soil without shrubs). Three main factors were found to determine the probability of sapling attack: sapling palatability, experimental microhabitat and plot. Palatable saplings (maples) were browsed much more than unpalatable ones (pines). The degree of protection provided by the shrub proved greater as its palatability decreased with respect to sapling palatability, the unpalatable spiny shrub being the safest microhabitat for palatable saplings and bare soil for unpalatable ones. The differences found in number of attacked saplings between plots may be attributable to differences in herbivore pressure. The community context in which interaction takes place, namely the characteristics of the neighbours and the intensity of herbivore pressure, are determining factors for understanding and predicting the damage undergone by a target plant species. The mechanism that best explains these results is associational avoidance of saplings that grow near to unpalatable shrubs. It is necessary to introduce this neighbour effect in theoretical models and food-web approaches that analyse the plant-herbivore relationships, since it can strongly determine not only the intensity of the interaction, but also the spatial distribution and diversity of the plant community.

While investigating biodiversity patterns on different spatial scales, ecological processes determining these patterns have been rarely analysed. Flower visitation by bees is an important ecological process that is related to floral resource availability. However, little is known about whether responses of bee communities to floral resource availability change at different spatial scales. We studied density and species richness of flower-visiting bees in relation to floral resource availability, provided by coffee, in traditional agroforestry systems on a field, shrub, and branch scale. On a field scale, mean bee density per shrub increased with decreasing proportion of flowering coffee shrubs per site, showing a dilution effect. Conversely, on shrub and branch scales bee density per shrub, or shrub part, increased with increasing number of inflorescences, showing a concentration effect. Additionally, bee density per shrub was higher on those that were only partly, rather than totally surrounded by other flowering coffee shrubs. Species richness of flower-visiting bees was positively affected by high resource availability on a shrub and a branch scale, expressed by a high number of inflorescences, but at the field scale the proportion of flowering shrubs per site did not have any effect on species richness. Our results show contrasting responses of the community of flower-visiting bees to floral resource availability, depending on the spatial scale considered. We conclude that patterns of flower-visiting bee communities of only one spatial scale can not be generalized, since the number of pollinators may be limited on a field scale, but not on smaller scales.

Herbivores can have indirect effects on local nutrient availability if their direct effects on plants lead to changes in the amount or chemical composition of litter reaching the soil surface. Using two exclosure experiments, we evaluated this possibility for blacktailed deer (Odocoileus hemionus columbianus ) feeding on silver bush lupine (Lupinus chamissonis ) in a coastal dune system in northern California. Our first experiment assessed the effects of deer herbivory on 360 lupines that were tracked from newly germinated seedlings in 1996 and 1997 until 2000, when most surviving individuals had reached reproductive age. Results from this experiment showed that browsing by deer significantly reduced growth rates and seed production of lupines and delayed the onset of reproductive maturity. Although deer had no effect on the density of litter accumulating underneath shrubs (g m 2 ), browsing significantly decreased C:N ratios of leaf tissue, due primarily to increased nitrogen content of leaves. Deer browsing also caused significantly increased net nitrogen mineralization rates in the soil under shrubs, although pools of ammonium and nitrate were uninfluenced. The second experiment examined 72 established bush lupines from 1997 to 2000, and showed that deer browsing significantly decreased seed production but had no effect on shrub growth. We also detected trends for browsed shrubs to have reduced pools of ammonium and nitrate underneath their canopies, although no such patterns emerged for nitrogen mineralization rates. Collectively, these data suggest that changes in litter quality, more so than litter quantity, contributed to the indirect effects of deer herbivory on local nitrogen dynamics. We hypothesize that deer browsing induced chemical changes in lupine plant tissue, which increased the nitrogen content of litter reaching the soil surface and subsequently increased rates of nitrogen mineralization.

Arid and semi-arid ecosystems often exhibit diverse plant growth forms in water-limited environments, but it is unclear whether resource competition (interference) is actually important in structuring communities. We chose a diverse Chihuahuan desert shrubland to examine the response of the plant community to experimental removals of selected perennial plant species or groups of species. Four treatments involved the removal of all individuals of all species of a single functional group (functional group removals: shrub removal, succulent removal, subshrub removal, perennial grass removal). Three other treatments involved removing species within functional groups. These seven treatments plus a control (no plants removed) were replicated six times each in 25 x25 m experimental plots, in summer 1995. Permanent belt transects were surveyed for number and sizes of all vascular plants in spring and fall in 1997, 1999, 2000, and 2001. Those plots from which the dominant shrub, Larrea tridentata, was removed had not recovered in total plant cover or volume by 2001, but cover and volume in all other treatments were similar to those in control plots. Relatively few species demonstrated a positive response to the removal of other species or functional groups. The perennial grass group and forbs were the most responsive; perennial grass cover increased in the shrub removal treatment relative to the control but treatment differences diminished after dry growing seasons in 2000 and 2001. Results over the first five years suggest that either environmental conditions or intrinsic biological characteristics limit the ability of most plant species to respond to the removal of substantial fractions of community biomass and composition in the short term. Such slow response by both dominant and less abundant components of the community has implications for the recovery of semi-arid systems after human disturbance or other events leading to the reduction of biological diversity.

Recent studies of ungulates have revealed that selective foraging seems to be an important mechanism by which they can affect the structure and species composition of the plant community, and thus quantity (dry mass) and quality (chemical composition) of litter available for decomposers. Such changes in litter production may be especially important in N-limited systems like boreal forests. We chose moose (Alces alces) as study species to investigate this mechanism. Moose browse mainly in the tree and shrub layers year round, and because of their wide distribution and often high population densities, they can have a significant effect on litter production of trees and shrubs in Swedish boreal forests. The effects of herbivores may also vary along productivity gradients. We therefore simulated browsing and urine and fecal deposition corresponding to 4 different moose densities in exclosures along a pre-existing forest productivity gradient. Both litter quantity (g dry mass per m 2 and year) and contributions of C and N (g dry mass per m 2 and year) decreased with increasing level of simulated moose density. High moose densities over extended time can therefore reduce N contributions to soil and therefore eventually reduce site productivity in Swedish boreal forests. This effect of moose was mainly a result of decreased litter quantity, because contradictory to studies from North America, litter quality (C:N ratio and N contribution per mass unit of litter) was not affected by level of simulated moose density.

A widely-used description of vegetation response to fire is that species can be clearly classified as sprouters or non-sprouters. We aimed to assess: (1) how well this dichotomous classification (sprouter/non-sprouter) described the responses of a semi-arid flora to experimental disturbance; (2) how similar were sprouting responses to treatments mimicking intense herbivory and fire; (3) how well easily-measured traits could predict sprouting. Sprouting was assessed for 45 species from a range of growth forms (grasses, forbs, sub-shrubs, woody shrubs and trees) from semi-arid south-eastern Australia. We used two treatments: clipping at stem base, and clipping followed by burning with a blowtorch. A dichotomy accounted for > 60% of deviance explained by species identity. Models with three or four groups were not substantially better. The dichotomy was not between 0% and 100% sprouting, rather between 'weak' and 'strong' sprouters. Probabilities of sprouting for weak sprouters were 23% after clipping and 6% after burning, while strong sprouters had sprouting probabilities of 90% after clipping and 79% after burning. While sprouting varied in space and time, the dichotomy was robust to this variation. Sprouting ability increased with size in most of the species with variable sprouting. Sprouting was partially related to growth form; grasses sprouted strongly, chenopods weakly, and forb and woody species covered the range of sprouting. Strong sprouters were likely to have more stems per plant, greater basal area, shorter potential height and deeper buds than weak sprouters. A hierarchical model that used growth form and then stems-per-plant provided a simple, robust predictor of sprouting. Four-fifths of species responded consistently to clipping and burning while one-fifth of species were strong sprouters after clipping but weak sprouters after burning. Burning reduced sprouting most in intermediate sprouters. Differences between sprouting after clipping and burning reflected increased intensity and were related to the depth of buds below ground.

Diversity in seedling responses to combined shade and drought can drive species niche differences, and thus natural forest and scrub establishment dynamics and diversity. However, inherent trade-offs between shade tolerance and drought tolerance, as hypothesized in the literature, would constrain potential niche differentiation. For thirteen species of European trees and shrubs the impacts were determined of moderate and extreme drought on the growth and survival of first-year seedlings in a given soil volume, in irradiances in the range typical for temperate forest and scrub understories (3% daylight) and large gaps (30% daylight). Pots were watered differentially each second day to equalize soil water content across species and irradiances. Comparisons within and across species supported independent tolerances of shade and drought rather than trade-offs. For all species, drought reduced relative growth rate in dry mass by the same proportion in 3% daylight and in 30% daylight. Consequently, drought generally reduced final dry mass significantly more strongly in 30% daylight than in 3% daylight. Extreme drought led to earlier mortality in 30% daylight than in 3% daylight for nine of the eleven species tested, to earlier mortality in 3% daylight for one species, and to equal mortality rate in both irradiances for one species. For each species, growth-based shade tolerance was quantified as the ratio of absolute growth in low irradiance to that in high irradiance, and growth-based drought tolerance as the ratio of absolute growth in low water supply to that in high water supply. Across species, growth-based shade tolerance varied ≈9-fold, and growth-based drought tolerance ≈2-fold; species’ tolerances correlated respectively with indices of field establishment in shade and drought. Growth-based drought tolerance correlated with survival time in extreme drought. Experimentally quantified shade and drought tolerances varied independently for the 13 species tested, indicating the potential for extensive species niche differentiation in combinations of irradiance and water supply.

Compensatory population dynamics among species stabilise aggregate community variables. Inter-specific competition is thought to be stabilising as it promotes asynchrony among populations. However, we know little about other inter-specific interactions, such as facilitation and granivory. Such interactions are also likely to influence population synchrony and community stability, especially in harsh environments where they are thought to have relatively strong effects in plant communities. We use a manipulative experiment to test the effects of granivores (harvester ants) and nurse plants (dwarf shrubs) on annual plant community dynamics in the Negev desert, Israel. We present evidence for weak and inconsistent effects of harvester ants on plant abundance and on population and community stability. By contrast, we show that annual communities under shrubs were more species rich, had higher plant density and were temporally less variable than communities in the inter-shrub matrix. Species richness and plant abundance were also more resistant to drought in the shrub under-storey compared with the inter-shrub matrix, although population dynamics in both patch types were synchronised. Hence, we show that inter-specific interactions other than competition affect community stability, and that hypothesised mechanisms linking compensatory dynamics and community stability may not operate to the same extent in arid plant communities.

Browsing by large vertebrates has been a major force in the evolution of terrestrial plants but Holocene extinctions of the browsers have left a legacy of broken biotic partnerships. Ratite birds were the largest herbivores in several regions, such as the moas of New Zealand. Many woody plants there have a distinct form of branching, described as “divaricate”, with thin, wide angled, branches intertwining to form a tangled canopy. Divaricate branching has been interpreted as a form of protection against climate extremes or as an anachronistic defense against the extinct moas. Here we report the first experimental evidence that many of these plants are defended against extant ratite browsers. In feeding experiments on two tree species with different (heteroblastic) juvenile and adult branch morphology, emus and ostriches obtained adequate feeding rates from adult shoots but sub-maintenance feeding rates from juvenile shoots with the ratite-resistant traits. Divaricate juvenile shoots suffered 30–70% less biomass removal to the birds than adult shoots. Ratites browse by a distinctive clamping and tugging action. Structural defence traits that exploit the limitations of this feeding mode include narrow, strong, elastic branches that resist being torn off, wide branching angle (“divaricate”) that makes shoots difficult to swallow, and small, widely spaced leaves. This novel plant architectural defence has developed in at least 20% of the native woody flora of New Zealand, including 10 heteroblastic tree species that exhibit the ratite-resistant strategy until they reach ca 3 metres height. It is also a major axis of variation amongst homoblastic woody shrub species. The defences are useless against mammalian browsers that shear shoots, contributing to marked decreases in the abundances of ratite-resistant species in New Zealand after the introduction of mammals.

In arctic Alaska, researchers have manipulated air temperature, light availability, and soil nutrient availability in several tundra communities over the past two decades. These communities responded quite differently to the same manipulations, and species responded individualistically within communities and among sites. For example, moist acidic tundra is primarily nitrogen (N)-limited, whereas wet sedge tundra is primarily phosphorus (P)-limited, and the magnitude of growth responses varies across sites within communities. Here we report results of four years of manipulated nutrients (N and/or P) and/or air temperature in an understudied, diverse plant community, moist non-acidic tussock tundra, in northern Alaska. Our goals were to determine which factors limit above-ground net primary productivity (ANPP) and biomass, how community composition changes may affect ecosystem attributes, and to compare these results with those from other communities to determine their generality. Although relative abundance of functional groups shifted in several treatments, the only significant change in community-level ANPP and biomass occurred in plots that received both N and P, driven by an increase in graminoid biomass and production resulting from a positive effect of adding N. There was no difference in community biomass among any other treatments; however, some growth forms and individual species did respond. After four years no one species has come to dominate the treatment plots and species richness has not changed. These results are similar to studies in dry heath, wet sedge, and moist acidic tundra where community biomass had the greatest response to both N and P and warming results were more subtle. Unlike in moist acidic tundra where shrub biomass increased markedly with fertilization, our results suggest that in non-acidic tundra carbon sequestration in plant biomass will not increase substantially under increased soil nutrient conditions because of the lack of overstory shrub species.

Understanding positive interactions among species is integral to predicting the outcomes of community-level dynamics. Few studies have explored intraspecific variation in positive associations. We investigated facilitation between two coastal dune shrubs in California, Baccharis pilularis and the nitrogen-fixing species, Lupinus arboreus. L., arboreus seedlings and adults were non-randomly distributed, occurring most frequently in association with B. pilularis. The relationship depended on the genotype of B. pilularis; only the prostrate architectural form of this species was positively associated with L. arboieus. To evaluate the nature of this association, we assessed the influence of three microhabitat types (prostrate B. pilularis, Ammophila arenaria (an introduced dune grass), and exposed sand) on four life history stages of L. arboreus. Prostrate B. pilularis benefited L. arboreus seedling emergence, survival, and growth but had no effect on post-dispersal seed predation or adult establishment. Enhanced resistances to granivory by rodents or to seedling predation by voles were not important mechanisms underlying facilitation. In addition, an experiment manipulating herbivory by cutworms found no effects of microhabitat on susceptibility to damage. Evidence supports the hypothesis that the prostrate form of B. pilularis facilitates L. arboreus germination, seedling growth, and establishment as compared to exposed sand and dune grass. Furthermore, in combination with prior research, the results suggest that by facilitating an important nitrogen-fixer, B. pilularis may have effects that cascade to other members of the coastal plant community.

We examined regrowth architecture of 4 species of savanna shrubs following 4 levels of defoliation. Defoliation had little effect on the regrowth architecture of honey mesquite (Prosopis glandulosa), which is rarely browsed by mammalian herbivores. The 3 acacia species (Acacia berlandieri, A. greggii, A. schaffneri) responded to defoliation by increasing leaf and spine density on the regrowth branches, but spine length and branching architecture remained unchanged. Only A. greggii, which is a preferred food plant of many browsers, exhibited an increase in the number and length of current annual growth branches in response to defoliation. The changes in plant architecture due to defoliation had little effect on the subsequent feeding behavior of captive white-tailed deer (Odocoileus virginianus). Food intake rate of the deer was most strongly related to internode distance, a parameter not significantly altered by defoliation. This suggests that the architectural responses of these shrubs to defoliation may not provide increased defense against browsing by co-evolved mammals.

12 pages, 10 tables, 2 figures. We thank Georgina Alvarez for the organizational support in our fieldtrips to Chafarinas. T. Gomez, M. Igual, J. Charco, and P. Robles gave field assistance. R. Lopez, J.M. Alegre, J. Cara and E. Gutierrez carried out plant and soil chemical analyses. D. G. Sobey kindly provided some key references. J.M. Murillo and L. Sack made valuable comments on an earlier version of the manuscript.

The spatial pattern of defoliation by mammalian herbivores across vegetation mosaics bas been frequently discussed, but rarely spatially quantified. Here we considered the role of plant-herbivore interactions in determining the spatial distribution of shrub defoliation by a large mammalian herbivore across a grass-shrub mosaic. We investigated the spatial pattern of heather defoliation by sheep in heather-grass mosaics. Heather-grass mosaics are two-phased vegetation mosaics, in which a spatially localized plant community (grass) fulfils nutritional needs, whilst a spatially extensive plant community (heather) meets energy requirements but is nutritionally marginal. We used a spatial analysis method, originating from human geography, to show that heather defoliation was not spread across the mosaic homogeneously, but that the spatial pattern was determined by geometric characteristics of the mosaic, grazing intensity, and the contrast between preferred and less preferred communities. The spatial analysis method proved to be a powerful tool to describe the spatial pattern of shrub defoliation. Applications of the method are explored and the implications of the spatial distribution of shrub defoliation are discussed.

Heavy grazing and trampling by reindeer increase nutrient cycling and primary production in areas where grasslands have replaced shrub and moss tundra. One way in which herbivores can affect nutrient cycling is through changing the litter decomposition processes. We studied the effect of herbivory on litter decomposition rate by reciprocal transplantation of litter between lightly grazed and heavily grazed areas, using the litterbag technique. We used litter from two of the most common species on the lightly grazed side, Betula nana and Empetrum nigrum, and two of the most common species on the heavily grazed side, Carex bigelowii and Deschampsia flexuosa. We found that herbivory improved litter quality by favouring species with easily decomposed litter. However, herbivory also improved litter quality by increasing the nitrogen content and lowering the C/N ratio of each species. Decomposition rates even correlated with the abundance of the plant category in question. Shrub litter decomposed faster in the lightly grazed area where shrubs were common, and graminoid litter decomposed faster in the heavily grazed area where graminoids were common. These results indicate that the decomposer micro-organisms are adapted to the most common litter types. This study shows that detailed information about the effect of herbivory on litter quality is important to understand differences between the short-term and long-term effects of herbivory on nutrient cycling and primary production.

Potential factors that trigger the onset of reproduction in shoot modules were studied in the dwarf shrub Rhododendron lapponicum in subarctic Swedish Lapland. The following questions were addressed: (1) Can the large size effect that commonly is found on onset of reproduction be explained as an effect of the amount of nitrogen and/or phosphorus accumulated? (2) Is there an effect of age when controlling for size and amount of nutrient reserves? (3) Does the relationship between size or nutrient reserves and onset of reproduction vary between high- and low-altitude sites? Shoots from two different sites, a subalpine heath (345 m altitude) and an alpine heath (900 m), were compared. The reproductive status of the shoots (non-reproductive or with an inflorescence bud) was related to shoot age, size (shoot leaf area), and nitrogen and phosphorus concentration by means of logistic regression analysis. The analysis indicates that an index of shoot nitrogen pool size was the most important factor; however, we could not strictly differentiate between the effect of this index and that of shoot leaf area and nitrogen concentration. A large shoot leaf area and nitrogen concentration (i.e. a large nitrogen pool index) increases the likelihood of turning reproductive. When controlling for shoot leaf area and nitrogen concentration a negative effect of increasing age was found. We suggest that this age effect may be related to a declining shoot resource economy with increasing age. Triggering of reproduction in R. lapponicum is thus complex and all factors included in our analysis appear to be involved. Shoots at the high-altitude site were more likely to enter the reproductive phase than shoots with a similar leaf area at low altitude. The only factor for which we found support for the hypothesis about decreasing importance of nutrients with increasing altitude was for phosphorus concentration. The mean age at onset of reproduction did not differ between altitudes.

A combination of competition and facilitation effects operating simultaneously among plant species appears to be the rule in nature, where these effects change along productivity gradients often in a non-proportional manner. We investigated changes in competition and facilitation between a leguminous shrub, Retama sphaerocarpa, and its associate understorey species along an environmental gradient in semi-arid southeast Spain. Our results show a change in the net balance of the interaction between the shrub and several of its associated species, from clearly positive in the water-stressed, infertile environment to neutral or even negative in the more fertile habitat. There was a weakening of facilitation along the fertility gradient as a consequence of improved abiotic conditions. Competition was the most intense for below-ground resources in the less fertile environment while total competition tended to increase towards the more productive end of the gradient. Changes in the balance of the interaction between and among different plant species along the gradient of stress were caused by a decline in facilitation rather than by a change in competition. As both competition intensity and facilitation change along gradients of resource availability, plant interactions are best viewed as dynamic relationships, the outcome of which depends on abiotic conditions.

High mortality of plants growing in dense monospecific stands (i.e. self-thinning) usually results from intense intraspecific competition. However, inconspicuous below-ground insect herbivory might be a potent but overlooked source of mortality within dense stands of plants, particularly if crowding limits a plant's ability to compensate for herbivore damage. Here I ask how high conspecific density influences a plant's ability to cope with heavy below-ground insect herbivory. I manipulated conspecific density and exposure to an abundant root-borer, the ghost moth (Hepialus californicus), and examined the impacts on the fecundity, growth, and survival of bush lupine (Lupinus arboreus), a fast-growing shrub that grows in dense monospecific stands in coastal grasslands, Both herbivory and intraspecific competition affected seed production, size, and mortality of bush lupine over the two years of the experiment. Plants consistently produced fewer seeds when growing at high versus low density and ghost moth herbivory also significantly reduced seed production. The negative effects of herbivory on plant fecundity were similar, regardless of plant density. In contrast, plant survival was affected by both competition, herbivory, and the interaction of these factors. In high density plots, plant survival was uniformly low (averaging 0.45-0.50); plants exposed to herbivores died from heavy herbivory, and plants protected from herbivores died due to intense intraspecific competition that compensated for losses due to herbivory. In low density plots, ghost moth herbivory similarly reduced lupine survival, from an average survival probability of 0.94 in plots protected from these herbivores to 0.55 in plots exposed to herbivory, Thus, results show that regardless of plant density, below-ground herbivory can be a potent source of mortality.