The responses of a community to disturbance: The importance of successional age and species' life histories

Summary The responses of different successional stages of a temperate intertidal algal community to disturbance were investigated with a field experiment. The experiment was conducted in a low intertidal boulder field in southern California. In this habitat, the top surfaces of boulders are covered with algae. The composition of the assemblage on any particular boulder depends on the length of time since it was last overturned by wave action. When a boulder is overturned, the algae on what was formerly the top surface, are killed in whole or part by a combination of sea urchin grazing, anoxia, light levels below compensation intensity, and mechanical damage caused by crushing or abrasion. The length of time that a boulder remains overturned and the local abundance of sea urchins determines the intensity of the disturbance. When the boulder is righted, recolonization begins either by vegetative regrowth of survivors and/or by spores from outside.Using a three-factorial design, this natural form of disturbance was experimentally mimicked and the responses of three different successional stages of the algal community monitored. Boulders in each successional category were overturned for periods of 17, 27 and 54 days in areas with and without sea urchins, then righted. Two aspects of community response to perturbation were evaluated. These were (1) the assemblage's ability to resist change and (2) its ability, if altered, to adjust to some semblance of its original state. The resistance of each assemblage and of its component species to change was measured by the percent decrease in algal cover and by the decline in percent similarity of the community to its original composition. The recovery rate of each assemblage and of the cover lost by each species during the first 35 days following a disturbance was measured by the rate of increase in percent similarity to the original composition and the percent reestablishment of lost cover.The experimental evidence demonstrates that the successional stages of the producer level of an intertidal algal community differ significantly in their responses to disturbance. Early successional communities suffer more damage from a given level of perturbation but recover more quickly than either middle or late successional communities. Damage to any particular assemblage of algae, irrespective of successional age, is more extensive and recovery slower, the longer the boulder is overturned and/or sea urchins are present. Several thresholds in these responses were also identified.Differences in community responses and non-linearities in these responses were attributable to the life history characteristics of the component species rather than emergent properties of the assemblage. These characteristics have evolved in response to a variety of recurrent natural disturbances. This interpretation is in agreement with recent critical reevaluations of the trends and mechanisms of successional change in natural communities.     

Predicting plant responses to mycorrhizae: integrating evolutionary history and plant traits

We assessed whether (1) arbuscular mycorrhizal colonization of roots (RC) and/or plant responses to arbuscular mycorrhizae (MR) vary with plant phylogeny and (2) MR and RC can be more accurately predicted with a phylogenetic predictor relative to a null model and models with plant trait and taxonomic predictors. In a previous study, MR and RC of 95 grassland species were measured. We constructed a phylogeny for these species and found it explained variation in MR and RC. Next, we used multiple regressions to identify the models that most accurately predicted plant MR. Models including either phylogenetic or phenotypic and taxonomic information similarly improved our ability to predict MR relative to a null model. Our study illustrates the complex evolutionary associations among species and constraints of using phylogenetic information, relative to plant traits, to predict how a plant species will interact with AMF.     

Contribution of physiological and morphological plant traits to a species' competitive ability at high and low nitrogen supply

Why do inherently fast-growing species from productive habitats generally have a higher rate of biomass production in short-term low-nitrogen experiments than slow-growing species from unproductive habitats, whereas the opposite is found in long-term experiments? Is this mainly due to inherent differences in biomass allocation, leaf characteristics or the plants' physiology? To analyse these questions we grew five monocotyledonous species from productive and unproductive habitats in a climate chamber at both high and low nitrogen supply. Nitrate was supplied exponentially, enabling us to compare inherent differences in morphological and physiological traits between the species, without any interference due to differences in the species' ability to take up nutrients. At high nitrogen supply, we found major inherent differences in specific leaf area and nitrogen productivity, i.e. daily biomass increment per unit plant nitrogen, where-as there were only small differences in net assimilation rate, i.e. daily biomass increment per unit leaf area, and biomass partitioning. We propose that the higher specific leaf area and nitrogen productivity of inherently fast-growing species are the key factors explaining their high abundance in productive habitats compared with inherently slow-growing ones. At low nitrogen supply, the net assimilation rate was decreased to a similar extent for all species, compared with that at high nitrogen supply. The nitrogen productivity of the inherentlyfast-growing species decreased with decreasing nitrogen supply, whereas that of the inherently slow-growing species remained constant. There were no inherent differences in nitrogen productivity in this treatment. At this low nitrogen supply, the inherently fast-growing species invested relatively more biomass in their roots that the slow-growing ones did. The inherently fast-growing species still had a higher specific leaf area at low nitrogen supply, but the difference between species was less than that at high nitrogen supply. Based on the present results and our optimization model for carbon and nitrogen allocation (Van der Werf et al. 1993a), we propose that the relatively large investment in root biomass of fast-growing species is the key factor explaining their higher biomass production in short-term experiments. We also propose that in the long run the competitive ability of the slow-growing species will increase due to a lower turnover rate of biomass. It is concluded that the plant's physiology (net assimilation rate and nitrogen productivity), only plays a minor role in the species' competitive ability in low-nitrogen environments.     

The epifauna of megaripples: species' adaptations and population responses to disturbance

Strong tidal currents flowing over mobile sediment give rise to migrating bedforms termed megaripples and sandwaves. For colonizing epifauna a megaripple field is a habitat subject to repeated disturbance as advancing megaripples bury all epifauna in their path. Eight epifaunal species occur in a megaripple field in tipper Spencer Gulf, South Australia. The stalked bryozoan Lanceopora obliqua is an r-strategist and occurs patchily in high densities. An ascidian Polycarpa pedunculata, probably a K-strategist, and a group of four species of articulated bryozoans, have much lower population densities which are predictable according to the frequency of disturbance. Other species occur in low, but variable densities. Some species show adaptations in morphology, behaviour or life history patterns which favour survival in this unusual environment.     

Combining plant and animal traits to assess community functional responses to disturbance

There is a general consensus that functional traits are reliable indicators of adaptation of organisms to particular environmental characteristics. In this study we relate the combined distributions of species traits of plants and animals to disturbance regimes in chestnut forests of southern Switzerland affected by regular winter fires. We used co‐inertia analysis for combining the trait response of 471 invertebrate species (117 001 individuals) and 81 plant species at 23 sites with different fire and cutting histories. Trait response was assessed by calculating the variation in weighted mean traits averaged over the communities and by using mean traits in multivariate analyses. The analysis showed a strong association between plant and animal traits under fire constraints (Monte‐Carlo test, p=0.0045). Plants and animal distributions show parallel trends in responses to fire which selects traits relating to persistence (ability to survive), resilience (ability to recover) and mobility. Warmth‐demanding insects, herbivores, flying carnivores and pollinators were associated with recent fires, as were annual, ruderal and light‐demanding plant species with long flowering duration. Small arthropods feeding on dead wood and those with narrow habitat requirements were associated with low fire frequency and unburnt sites, as were competitive plants with large seeds favoring moist sites. The spatial association between plant and animal traits reflected adaptations that promote survival in the disturbance regime, while the disturbance acts as an environmental filter on the distribution and assemblage of the trait values within communities. This combined analysis of plant and invertebrate traits distributions illustrates how community and ecosystem responses can be monitored and the results generalized across localities and disturbance types. Analyses of traits that cross trophic levels provide powerful and promising tools for validating management procedures and controlling ecosystem functions.     

浑善达克沙地中部斑块格局影响植物多样性及功能性状

景观格局与植物多样性之间的关系已被广泛研究,然而,斑块格局如何影响植物多样性,以及这些影响的边缘类型差异尚不清楚。为从植物功能性状角度揭示斑块格局影响植物多样性的机制,该研究采用不同边缘类型的斑块,利用两年调查的705个样方数据,结合遥感卫星图像解译和空间信息分析,根据植物群落的空间位置和斑块格局,按照向内、向外、向内成核和向外成核4种边缘类型将浑善达克沙地中部的斑块进行分类,采用Duncan新复极差法比较了4种边缘类型斑块格局与植物功能性状多样性关系的差异,应用Pearson相关系数和冗余分析(RDA)法量化了斑块格局对植物多样性的影响。结果发现,植物丰富度和多样性指数与斑块格局密切相关,这些关系存在着边缘类型的显著差异,表现出不同的功能性状。对于向内边缘,边缘指数与丰富度负相关,和C₄植物比例正相关;对于向外边缘,边缘指数与C₄植物比例正相关;对于向内成核边缘,边缘密度与多年生植物和虫媒植物比例正相关;对于向外成核边缘,边缘角度和指数与物种Shannon-Wiener指数和Simpson指数正相关,与Pielou指数、丰富度和动物传播植物...     

Geographical variation in species' population responses to changes in temperature and precipitation

Despite increasing concerns about the vulnerability of species'' populations to climate change, there has been little overall synthesis of how individual population responses to variation in climate differ between taxa, with trophic level or geographically. To address this, we extracted data from 132 long-term (greater than or equal to 20 years) studies of population responses to temperature and precipitation covering 236 animal and plant species across terrestrial and freshwater habitats. Our results identify likely geographical differences in the effects of climate change on populations and communities in line with macroecological theory. Temperature tended to have a greater overall impact on populations than precipitation, although the effects of increased precipitation varied strongly with latitude, being most positive at low latitudes. Population responses to increased temperature were generally positive, but did not vary significantly with latitude. Studies reporting significant climatic trends through time tended to show more negative effects of temperature and more positive effects of precipitation upon populations than other studies, indicating climate change has already impacted many populations. Most studies of climate change impacts on biodiversity have focused on temperature and are from middle to high northern latitudes. Our results suggest their findings may be less applicable to low latitudes.     

Legume species differ in the responses of their functional traits to plant diversity

Plants can respond to environmental impacts by variation in functional traits, thereby increasing their performance relative to neighbors. We hypothesized that trait adjustment should also occur in response to influences of the biotic environment, in particular different plant diversity of the community. We used 12 legume species as a model and assessed their variation in morphological, physiological, life-history and performance traits in experimental grasslands of different plant species (1, 2, 4, 8, 16 and 60) and functional group (1–4) numbers. Mean trait values and their variation in response to plant diversity varied among legume species and from trait to trait. The tall-growing Onobrychis viciifolia showed little trait variation in response to increasing plant diversity, whereas the species with shorter statures responded in apparently adaptive ways. The formation of longer shoots with elongated internodes, increased biomass allocation to supporting tissue at the cost of leaf mass, reduced branching, higher specific leaf areas and lower foliar δ¹³C values indicated increasing efforts for light acquisition in more diverse communities. Although leaf nitrogen concentrations and shoot biomass:nitrogen ratios were not affected by increasing plant diversity, foliar δ¹⁵N values of most legumes decreased and the application of the ¹⁵N natural abundance method suggested that they became more reliant on symbiotic N₂ fixation. Some species formed fewer inflorescences and delayed flowering with increasing community diversity. The observed variation in functional traits generally indicated strategies of legumes to optimize light and nutrient capturing, but they were largely species-dependent and only partly attributable to increasing canopy height and community biomass with increasing plant diversity. Thus, the analysis of individual plant species and their adjustment to growth conditions in communities of increasing plant diversity is essential to get a deeper insight into the mechanisms behind biodiversity–ecosystem functioning relationships.     

Plant responses to variable timing of aboveground clipping and belowground herbivory depend on plant age

Aims Plants use different types of responses such as tolerance and induced defense to mitigate the effects of herbivores. The direction and magnitude of both these plant responses can vary with plant age. However, most studies have focused on aboveground herbivory, whereas important feeding occurs belowground. Here, we tested the hypothesis that plant tolerance and defense following shoot damage or root herbivory depends on plant age.     

CO2‐mediated changes of plant traits and their effects on herbivores are determined by leaf age

1. Concentration of atmospheric CO₂ is predicted to double during the 21st century. However, quantitative effects of increased CO₂ levels on natural herbivore–plant interactions are still little understood. 2. In this study, we assess whether increased CO₂ quantitatively affects multiple defensive and nutritive traits in different leaf stages of cyanogenic wildtype lima bean plants (Phaseolus lunatus), and whether plant responses influence performance and choice behaviour of a natural insect herbivore, the Mexican bean beetle (Epilachna varivestis). 3. We cultivated lima bean plants in climate chambers at ambient, 500, 700, and 1000 ppm CO₂ and analysed cyanogenic precursor concentration (nitrogen‐based defence), total phenolics (carbon‐based defence), leaf mass per area (LMA; physical defence), and soluble proteins (nutritive parameter) of three defined leaf age groups. 4. In young leaves, cyanide concentration was the only parameter that quantitatively decreased in response to CO₂ treatments. In intermediate and mature leaves, cyanide and protein concentrations decreased while total phenolics and LMA increased. 5. Depending on leaf stage, CO₂‐mediated changes in leaf traits significantly affected larval performance and choice behaviour of adult beetles. We observed a complete shift from highest herbivore damage in mature leaves under natural CO₂ to highest damage of young leaves under elevated CO_2. Our study shows that leaf stage is an essential factor when considering CO₂‐mediated changes of plant defences against herbivores. Since in the long run preferred consumption of young leaves can strongly affect plant fitness, variable effects of elevated CO₂ on different leaf stages should receive highlighted attention in future research.     

Direct and indirect effects of pollinators and seed predators to selection on plant and floral traits

Although flowering traits are often assumed to be under strong selection by pollinators, significant variation in such traits remains the norm for most plant species. Thus, it is likely that the interactions among plants, mutualists, and other selective agents, such as antagonists, ultimately shape the evolution of floral and flowering traits. We examined the importance of pollination vs pre-dispersal seed predation to selection on plant and floral characters via female plant-reproductive success in Castilleja linariaefolia (Scrophulariaceae). C. linariaefolia is pollinated by hummingbirds and experiences high levels of pre-dispersal seed predation by plume moth and fly larvae in the Rocky Mountains of Colorado, USA, where this work was conducted. We first examined whether female reproduction in C. linariaefolia was limited by pollination. Supplemental pollination only marginally increased components of female reproduction, likely because seed predation masked, in part, the beneficial effects of pollen addition. In unmanipulated populations, we measured calyx length, flower production, and plant height and used path analysis combined with structural equation modeling to quantify their importance to relative seed set through pathways involving pollination vs seed predation. We found that the strength of selection on calyx length, flower production, and plant height was greater for seed predation pathways than for pollination pathways, and one character, calyx length, experienced opposing selection via pollination vs seed predation. These results suggest that the remarkable intraspecific variation in plant and floral characters exhibited by some flowering plants is likely the result of selection driven, at least in part, by pollinators in concert with antagonists, such as pre-dispersal seed predators. This work highlights the subtle but complex interactions that shape floral and vegetative design in natural ecosystems.     

A comparison of the responses of two tropical specialist herbivores to host plant patch size

Summary The effects of host plant patch size on the abundances of two specialist herbivores (the chrysomelid beetle, Acalymma innubum and the pentatomid bug, Piezosternum subulatum) were investigated in a natural forest community in the Virgin Islands. Abundances were compared early and late in the season in different sized patches of the cucurbit host plant (Cayaponia americana) growing in open habitat (with no surrounding plant community) and forest habitat (with diverse surrounding plant community). For both herbivore species, adult abundances per patch were positively correlated with patch leaf area, but there was a significant patch size effect (i.e., correlation between herbivore density per unit plant and patch leaf area) only for beetles in the forest habitat. Both herbivore species were significantly affected by surrounding plant diversity, but in opposite ways: beetles were more abundant in open patches whereas bugs were more abundant in forest patches. Relationships between abundance and patch size in open and forest patches changed through the season for both herbivore species. These changing abundance patterns are discussed with respect to (1) increases in the diversity of the plant community surrounding host plant patches, and (2) differences in herbivore movement patterns.     

Host patch traits have scale-dependent effects on diversity in a stickleback parasite metacommunity

Many metacommunities are distributed across habitat patches that are themselves aggregated into groups. Perhaps the clearest example of this nested metacommunity structure comes from multi-species parasite assemblages, which occupy individual hosts that are aggregated into host populations. At both spatial scales, we expect parasite community diversity in a given patch (either individual host or population) to depend on patch characteristics that affect colonization rates and species sorting. But, are these patch effects consistent across spatial scales? Or, do different processes govern the distribution of parasite community diversity among individual hosts, versus among host patches? To answer these questions, we document the distribution of parasite richness among host individuals and among populations in a metapopulation of threespine stickleback Gasterosteus aculeatus. We find some host traits (host size, gape width) are associated with increased parasite richness at both spatial scales. Other patch characteristics affect parasite richness only among individuals (sex), or among populations (lake size, lake area, elevation and population mean heterozygosity). These results demonstrate that some rules governing parasite richness in this metacommunity are shared across scales, while others are scale-specific.     

The effects of age at mating on female life-history traits in a seed beetle

Age at first reproduction is an important component of lifehistory across taxa and can ultimately affect fitness. Becausegenetic interests of males and females over reproductive decisionscommonly differ, theory predicts that conflict may arise overthe temporal distribution of matings. To determine the potentialfor such sexual conflict, we studied the direct costs and benefitsassociated with mating at different times for females, usingseed beetles (Acanthoscelides obtectus) as a model system. Virginfemales were resistant to male mating attempts at a very earlyage but subsequently reduced their resistance. Although we foundno difference in life span or mortality rates between femalesmated early in life and those mated later, females that matedearly in life suffered a 12% reduction in lifetime fecundity.Thus, there are direct costs associated with mating early inlife for females. Yet, males mate even with newly hatched females.We suggest that these data indicate a potential for sexual conflictover the timing of first mating and that female resistance tomating, at least in part, may represent a female strategy aimedat delaying mating to a later time in life.     

Forest fragmentation effects on patch occupancy and population viability of herbaceous plant species

Habitat fragmentation is one of the major threats to species diversity. In this review, we discuss how the genetic and demographic structure of fragmented populations of herbaceous forest plant species is affected by increased genetic drift and inbreeding, reduced mate availability, altered interactions with pollinators, and changed environmental conditions through edge effects. Reported changes in population genetic and demographic structure of fragmented plant populations have, however, not resulted in large-scale extinction of forest plants. The main reason for this is very likely the long-term persistence of small and isolated forest plant populations due to prolonged clonal growth and long generation times. Consequently, the persistence of small forest plant populations in a changing landscape may have resulted in an extinction debt, that is, in a distribution of forest plant species reflecting the historical landscape configuration rather than the present one. In some cases, fragmentation appears to affect ecosystem integrity rather than short-term population viability due to the opposition of different fragmentation-induced ecological effects. We finally discuss extinction and colonization dynamics of forest plant species at the regional scale and suggest that the use of the metapopulation concept, both because of its heuristic power and conservation applications, may be fruitful.