Stochastic and deterministic processes together determine alpine meadow plant community composition on the Tibetan Plateau

To predict the consequences of environmental change on the structure and composition of communities, it is necessary to also understand the regional drivers underlying the structuring of these communities. Here, we have taken a hypothesis-based approach to test the relative importance of niche versus neutral processes using niche overlap, species traits and population asynchrony in two crossed treatments of fertilization and grazing in an alpine meadow community. Our results suggested that the observed species biomass overlap was not significantly different between treatments of grazing, grazing × fertilization and grazer exclusion. In contrast, the species biomass overlap was higher than expected in fertilization treatments when grazers were excluded. On the one hand, we found no relationship between species traits and relative abundance in grazing, grazing × fertilization and grazer-exclusion treatments; on the other hand, mechanistic trait-based theory could be used to predict species relative abundance patterns in fertilization treatments when grazers were excluded. From grazing to fertilization, when grazers were excluded, there was a slight increase in species synchrony, which indicated that the complementary dynamic of species gradually changed from complete independence into synchronously fluctuating with increasing fertilization. Based on the above results, we concluded that stochastic and deterministic processes formed ends of a continuum from grazing to fertilization when grazers were excluded in an alpine meadow plant community, and the importance of niche differences between species in structuring grassland communities increased with increasing fertilization and decreased with grazing.     

The effects of fertilization on the trait-abundance relationships in a Tibetan alpine meadow community

Aims Comparisons of the trait–abundance relationships from various habitat types are critical for community ecology, which can offer us insights about the mechanisms underlying the local community assembly, such as the relative role of neutral vs. niche processes in shaping community structure. Here, we explored the responses of trait–abundance relationships to nitrogen (N), phosphorus (P) and potassium (K) fertilization in an alpine meadow.Methods Five fertilization treatments (an unfertilized control and additions of N, P, K and NPK respectively) were implemented using randomized block design in an alpine Tibetan meadow. Species relative abundance (SRA), plant above-ground biomass and species richness were measured in each plot. For 24 common species, we measured species functional traits: saturated height, specific leaf area (SLA) and leaf dry matter content (LDMC) in each treatment but seed size only in the unfertilized control. Standard major axis (SMA) regression and phylogenetically independent contrasts (PICs) analysis were used to analyse species trait–abundance relationships in response to different fertilization treatments.Important findings Positive correlations between SRA and saturated height were raised following N, P and NPK fertilizations, which indicated an increase in light competition in these plots. In P fertilized plots, SRA was also positively correlated with LDMC because tall grasses with a nutrients conservation strategy often have a relative competitive advantage in capturing limited light and soil nutrients. In K fertilized plots, neither the trait–abundance relationships nor above-ground biomass or species richness significantly differed from that in the control, which suggests that K was not a limiting resource in our study site. These significant correlations between species traits and relative abundance in fertilized treatment suggest that trait-based selection plays an important role in determining species abundance within local communities in alpine meadows.     

Plant functional and phylogenetic turnover correlate with climate and land use in the Western Swiss Alps

Aims Understanding the relative importance of historical and environmental processes in the structure and composition of communities is one of the longest quests in ecological research. Increasingly, researchers are relying on the functional and phylogenetic β-diversity of natural communities to provide concise explanations on the mechanistic basis of community assembly and the drivers of trait variation among species. The present study investigated how plant functional and phylogenetic β-diversity change along key environmental and spatial gradients in the Western Swiss Alps.Methods Using the quadratic diversity measure based on six functional traits—specific leaf area, leaf dry matter content, plant height, leaf carbon content, leaf nitrogen content and leaf carbon to nitrogen content alongside a species-resolved phylogenetic tree—we relate variations in climate, spatial geographic, land use and soil gradients to plant functional and phylogenetic turnover in mountain communities of the Western Swiss Alps.Important findings Our study highlights two main points. First, climate and land-use factors play an important role in mountain plant community turnover. Second, the overlap between plant functional and phylogenetic turnover along these gradients correlates with the low phylogenetic signal in traits, suggesting that in mountain landscapes, trait lability is likely an important factor in driving plant community assembly. Overall, we demonstrate the importance of climate and land-use factors in plant functional and phylogenetic community turnover and provide valuable complementary insights into understanding patterns of β-diversity along several ecological gradients.     

Aboveground resilience to species loss but belowground resistance to nitrogen addition in a montane plant community

Aims Decades of empirical work have demonstrated how dominant plant species and nitrogen fertilization can influence the structure and function of plant communities. More recent studies have examined the interplay between these factors, but few such studies use an explicit trait-based framework. In this study, we use an explicit trait-based approach to identify potential mechanisms for community-level responses and to test ecological niche theory.Methods We experimentally manipulated plant communities (control, ?dominant species, ?random biomass) and nitrogen (N) inputs (control, +organic N, +inorganic N) in a fully factorial design. We predicted that traits related to plants' ability to take up different forms of soil N would differ between dominant and subordinate species, resulting in interactive effects of dominant species loss and N fertilization on plant community structure and function. The study took place in a montane meadow in the Rocky Mountains, Colorado, USA.Important findings After four years, the plant community in removal plots converged toward a species composition whose leaf and root functional traits resembled those of the previously removed dominant species. Ecosystem productivity generally increased with N addition: soil carbon efflux was ~50% greater when either form of N was added, while inorganic N addition increased aboveground biomass production by ~60% relative to controls. The increase in production was mediated by increased average height, leaf mass:area ratio and leaf dry matter content in plant communities to which we added inorganic N. Contrary to our predictions, there were no interactive effects of N fertilization and dominant species loss on plant community structure or ecosystem function. The plant community composition in this study exhibited resistance to soil N addition and, given the functional convergence we observed, was resilient to species loss. Together, our results indicate that the ability of species to compensate functionally for species loss confers resilience and maintains diversity in montane meadow communities.     

Trait‐based community assembly of aquatic macrophytes along a water depth gradient in a freshwater lake

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Does niche overlap control relative abundance in French lacustrine fish communities? A new method incorporating functional traits

1. The mechanisms that structure biological communities hold the key to understanding ecosystem functioning and the maintenance of biodiversity. Patterns of species abundances have been proposed as a means of differentiation between niche-based and neutral processes, but abundance information alone cannot provide unequivocal discrimination. 2. We combined species niche information and species' relative abundances to test the effects of two opposing structuring mechanisms (environmental filtering and niche complementarity) on species' relative abundances in French lacustrine fish communities. The test involved a novel method comparing the abundance-weighted niche overlap within communities against that expected when relative abundances were randomized among species within the community. 3. Observed overlap was consistently significantly lower than expected at random for two (swimming ability and trophic status) of four primary niche axes across lakes of differing physical environments. Thus, for these niche axes, pairs of abundant species tended to have relatively low niche overlap, while rare species tended to have relatively high niche overlap with abundant species. 4. This suggests that niche complementarity may have acted to enhance ecosystem function and that it is important for species coexistence in these fish communities. The method used may be easily applied to any sort of biological community and thus may have considerable potential for determining the generality of niche complementarity effects on community structure.     

Convergent succession of plant communities is linked to species’ functional traits

Functional traits reflecting the resource economy and growth strategy of plants vary widely both within and among ecosystems. Theory suggests that trait variation within a community may determine the relative abundance of species, though this idea requires more empirical support.We set up a long-term succession experiment in a nutrient-poor wetland, planting seedlings of twelve fenland species in different relative abundances and absolute densities, thereby creating 24 communities. The biomass of these species and the soil water and nutrient status of the system were monitored over ten years. Using these data, we could relate the changing relative abundance of species to five traits – leaf dry matter content (LDMC), leaf nitrogen concentration (LNC), specific leaf area (SLA), relative growth rate (RGR), and seed mass (SM).The initial communities converged after ten years to a common dominance–diversity structure, with two species accounting for 82% of total biomass. Soil water and nutrient conditions remained largely constant. By the end of the experiment, community trait structure had changed so that species functional traits were significantly related to their relative abundance. The most abundant species had high LDMC and SM, but low RGR and SLA, and varied little in LNC, suggesting that investment in leaf structure and retention of nutrients were most important for species dominance under low nutrient conditions. Our results provide experimental evidence that dominance–subdominance structures in plant communities are governed by functional traits.     

沙质草地不同生活史植物的生物量分配对氮素和水分添加的响应

以氮素和水分(冬季增雪和夏季增雨)为控制因子, 开展相关田间控制实验, 分析不同功能群(以生活史为划分依据)尺度和群落尺度植物生物量分配格局对氮素和水分的响应, 得出以下结论: 1)一年生植物的繁殖生物量比重明显高于多年生植物, 而多年生植物种的叶/地上生物量比值显著高于一年生植物; 2)一年生植物对氮素和水分添加的响应剧烈, 氮添加耦合夏季增雨、氮添加耦合冬季增雪显著增加了一年生植物的繁殖生物量比重和叶生物量比重。多年生植物对氮素和水分添加的响应不敏感, 表现为多年生植物的各器官生物量分配格局对氮素添加和水分添加的响应不明显。3)氮素添加和水分处理改变了群落尺度生物量分配格局: 氮素添加耦合冬季增雪处理降低了群落植物的繁殖生物量比重和茎生物量比重, 提高了群落植物的叶生物量比重。4)冬季增雪和夏季增雨与氮素添加的交互作用对群落生物量分配格局的改变不同。夏季增雨耦合氮素添加处理下群落的茎生物量比重显著提高, 群落茎生物量分配的改变引起群落的垂直结构发生改变。冬季增雪氮素处理下群落的叶生物量比重增加, 但茎生物量比重增加不明显。冬季增雪也改变了群落的结构和功能。     

Trends in Biomass Fractionation in Wheat and Barley from Wild Ancestors to Modern Cultivars

Abstract: We tested the hypothesis that cultivar selection during the process of domestication in cereal plants led to a change in dry mass allocation, e.g., less root mass and more leaf mass or more leaf area per unit leaf mass. We divided 24 varieties of diploid, tetraploid and hexaploid winter wheat and two-rowed winter barley into three categories of domestication levels (wild species, old landraces and modern cultivars) and compared the patterns of dry matter fractionation at the time of anthesis under standardized outdoor growth conditions. In both cereals, total biomass per individual increased significantly with domestication level but, to our surprise, we found no significant change in dry matter investment between domestication levels: neither the dry mass fraction of leaves increased, nor was there a trend of reduced investment in stems and roots, contrary to what we expected. Specific leaf area (SLA) and leaf area ratio (LAR) of modern wheat and barley cultivars were significantly lower compared to wild varieties. Major differences in both cereals were of a purely morphological nature, namely a decrease in the number of stems and ears from wild species to domesticated varieties, along with more synchronous tiller development and therefore similar tiller size. Fertilizer increased total biomass in all domestication levels in both cereals, but influenced the dry matter fractionation only in barley. Tissue nitrogen concentration was unresponsive to both domestication and fertilization. The expected shift in functional traits, conventionally considered to determine plant growth, was not found. Indeed, dry matter fractionation among the major plant biomass components seems to be very conservative.     

刈割与施肥干扰下高寒草甸植物功能性状的构建模式

针对目前关于植物群落功能性状构建模式及其驱动因素存在的争议, 研究了33个物种10个功能性状的构建模式。研究结果显示: (1)在刈割-施肥复合梯度上, 这些功能性状主要表现为随机构建模式, 发生随机性和确定性构建的样方比例分别为82.7%和17.3%; (2)在10个功能性状中, 生长型、生活周期、单株地上干质量、叶面积和叶干质量5个功能性状为随机构建模式, 不受试验处理和群落特征(地上净初级生产力、刈割生物量损失、群落高度)变化的影响。植物倾斜度、繁殖方式、固氮性、株高和比叶面积5个功能性状的构建模式与试验处理或群落特征变化有关, 表现出趋同或趋异构建响应。其中, 植物倾斜度和比叶面积的构建模式仅受群落特征影响, 而固氮性、繁殖方式和株高3个性状的构建模式同时还受刈割或施肥处理的影响, 其构建模式因功能性状而异; (3)群落的生物量损失和地上净初级生产力是解释植物功能性状构建模式变化较理想的群落特征; (4)刈割和施肥处理对株高的构建模式具有相反效应, 而刈割和施肥的交互作用对其无显著影响。上述结果说明该研究群落植物功能性状的构建存在不同模式, 以随机构建模式为主, 确定性构建模式居次要地位。确定性构建模式与试验处理和群落特征变化有关, 而且是性状依赖的。相反选择力对趋同和趋异构建模式的平衡效应能引起功能性状发生随机构建。     

Plant Trait Variation along an Altitudinal Gradient in Mediterranean High Mountain Grasslands: Controlling the Species Turnover Effect

Assessing changes in plant functional traits along gradients is useful for understanding the assembly of communities and their response to global and local environmental drivers. However, these changes may reflect the effects of species composition (i.e. composition turnover), species abundance (i.e. species interaction), and intra-specific trait variability (i.e. species plasticity). In order to determine the relevance of the latter, trait variation can be assessed under minimal effects of composition turnover. Nine sampling sites were established along an altitudinal gradient in a Mediterranean high mountain grassland community with low composition turnover (Madrid, Spain; 1940 m–2419 m). Nine functional traits were also measured for ten individuals of around ten plant species at each site, for a total of eleven species across all sites. The relative importance of different sources of variability (within/between site and intra-/inter-specific functional diversity) and trait variation at species and community level along the considered gradients were explored. We found a weak individual species response to altitude and other environmental variables although in some cases, individuals were smaller and leaves were thicker at higher elevations. This lack of species response was most likely due to greater within- than between-site species variation. At the community level, inter-specific functional diversity was generally greater than the intra-specific component except for traits linked to leaf element content (leaf carbon content, leaf nitrogen content, δ¹³C and δ¹⁵N). Inter-specific functional diversity decreased with lower altitude for four leaf traits (specific leaf area, leaf dry matter content, δ¹³C and δ¹⁵N), suggesting trait convergence between species at lower elevations, where water shortage may have a stronger environmental filtering effect than colder temperatures at higher altitudes. Our results suggest that, within a vegetation type encompassing various environmental gradients, both, changes in species abundance and intra-specific trait variability adjust for the community functional response to environmental changes.     

The role of biomass allocation strategy in diversity loss due to fertilization

Proposed mechanisms for explaining biodiversity loss due to fertilization include interspecific competition and assemblage-level thinning. The interspecific competition hypothesis (ICH) assumes a link between population changes and species competitive ability, which is related to functional traits such as biomass allocation patterns. Based on a 2-year fertilization experiment in an alpine meadow on the Tibetan Plateau, we attempted to explore the relationships between individual and population responses. Individual response was measured by changes in plant biomass and biomass allocation, and population response was estimated by changes in species abundance. The results suggested that following fertilization (1) changes in individual biomass differ among species and functional groups, (2) reproductive allocation tends to decrease for all species whereas leaf allocation generally increases for grasses but decreases for forbs, (3) a strong positive correlation exists between species relative abundance change and individual biomass response, and (4) species relative abundance change has a positive correlation with leaf allocation response, a negative correlation with stem allocation response, and no significant correlation with reproductive allocation response. We conclude that the individual biomass responses and biomass allocation strategy can partly explain diversity loss due to fertilization, a result consistent with the ICH.     

Trait‐abundance relation in response to nutrient addition in a Tibetan alpine meadow: The importance of species trade‐off in resource conservation and acquisition

In competition‐dominated communities, traits promoting resource conservation and competitive ability are expected to have an important influence on species relative abundance (SRA). Yet, few studies have tested the trait‐abundance relations in the line of species trade‐off in resource conservation versus acquisition, indicating by multiple traits coordination. We measured SRA and key functional traits involving leaf economic spectrum (SLA, specific leaf area; LDMC, leaf dry matter content; LCC, leaf carbon concentration; LNC, leaf nitrogen concentration; LPC, leaf phosphorus concentration; Hs, mature height) for ten common species in all plots subjected to addition of nitrogen fertilizer (N), phosphorus fertilizer (P), or both of them (NP) in a Tibetan alpine meadow. We test whether SRA is positively related with traits promoting plant resource conservation, while negatively correlated with traits promoting plant growth and resource acquisition. We found that species were primarily differentiated along a trade‐off axis involving traits promoting nutrient acquisition and fast growth (e.g., LPC and SLA) versus traits promoting resource conservation and competition ability (e.g., large LDMC). We further found that SRA was positively correlated with plant height, LDMC, and LCC, but negatively associated with SLA and leaf nutrient concentration irrespective of fertilization. A stronger positive height‐SRA was found in NP‐fertilized plots than in other plots, while negative correlations between SRA and SLA and LPC were found in N or P fertilized plots. The results indicate that species trade‐off in nutrient acquisition and resource conservation was a key driver of SRA in competition‐dominated communities following fertilization, with the linkage between SRA and traits depending on plant competition for specific soil nutrient and/or light availability. The results highlight the importance of competitive exclusion in plant community assembly following fertilization and suggest that abundant species in local communities become dominated at expense of growth while infrequent species hold an advantage in fast growth and dispersals to neighbor meta‐communities.     

Processes of community assembly in an environmentally heterogeneous,high biodiversity region

Despite decades of study, the relative importance of niche‐based versus neutral processes in community assembly remains largely ambiguous. Recent work suggests niche‐based processes are more easily detectable at coarser spatial scales, while neutrality dominates at finer scales. Analyses of functional traits with multi‐year multi‐site biodiversity inventories may provide deeper insights into assembly processes and the effects of spatial scale. We examined associations between community composition, species functional traits, and environmental conditions for plant communities in the Kouga‐Baviaanskloof region, an area within South Africa's Cape Floristic Region (CFR) containing high α and β diversity. This region contains strong climatic gradients and topographic heterogeneity, and is comprised of distinct vegetation classes with varying fire histories, making it an ideal location to assess the role of niche‐based environmental filtering on community composition by examining how traits vary with environment. We combined functional trait measurements for over 300 species with observations from vegetation surveys carried out in 1991/1992 and repeated in 2011/2012. We applied redundancy analysis, quantile regression, and null model tests to examine trends in species turnover and functional traits along environmental gradients in space and through time. Functional trait values were weakly associated with most spatial environmental gradients and only showed trends with respect to vegetation class and time since fire. However, survey plots showed greater compositional and functional stability through time than expected based on null models. Taken together, we found clear evidence for functional distinctions between vegetation classes, suggesting strong environmental filtering at this scale, most likely driven by fire dynamics. In contrast, there was little evidence of filtering effects along environmental gradients within vegetation classes, suggesting that assembly processes are largely neutral at this scale, likely the result of very high functional redundancy among species in the regional species pool.     

Quantifying the role of deterministic assembly and stochastic drift in a natural community of Arctic mosses

The interpretation of natural plant communities frequently invokes species‐sorting controlled by niche differences along spatial environmental gradients. This process of niche structuring can be explained by reference to functional traits, which provide a mechanistic explanation for community structure. In contrast, models explaining species coexistence obviate the limiting effect of niche difference, by invoking processes which cause species‐level drift, e.g. demographic stochasticity. This paper investigates a simple habitat with strong gradients (moss communities in a patterned arctic wetland) to identify signature‐patterns under‐pinning the relative importance of deterministic assembly and stochastic drift in a natural community. First, ordination analysis was used to confirm community composition structured by a range of nine carefully selected functional traits. Second, to determine whether traits explaining community composition might also explain species richness, local species richness (sR) was compared to (1) observed trait diversity and (2) expected trait diversity based on permutation tests, which are used to simulate null community assembly for different values of sR. Traits explaining species composition, consistent with deterministic niche structuring, do not appear to maintain sR. This surprising result was explained by decomposing the community into individual pair‐wise comparisons, i.e. species niche‐differences and association (χ²). Results support deterministic processes via the sorting of species with similar and contrasting niches, at opposite ends of a composite environmental gradient. Nevertheless, stochastic drift is apparent in the random structure of a majority of pair‐wise associations; in addition, a species’ abundance was in general not related to environmental distance from response‐optima. We suggest therefore that spatial pattern in the moss community is a balance between deterministic forces with respect to species traits and controlling environmental gradients, and stochastic drift, which weakens this deterministic structure.     

Taxa-area relationship and neutral dynamics influence the diversity of fungal communities on senesced tree leaves

This study utilized individual senesced sugar maple and beech leaves as natural sampling units within which to quantify saprotrophic fungal diversity. Quantifying communities in individual leaves allowed us to determine if fungi display a classic taxa–area relationship (species richness increasing with area). We found a significant taxa–area relationship for sugar maple leaves, but not beech leaves, consistent with Wright's species‐energy theory. This suggests that energy availability as affected plant biochemistry is a key factor regulating the scaling relationships of fungal diversity. We also compared taxa rank abundance distributions to models associated with niche or neutral theories of community assembly, and tested the influence of leaf type as an environmental niche factor controlling fungal community composition. Among rank abundance distribution models, the zero‐sum model derived from neutral theory showed the best fit to our data. Leaf type explained only 5% of the variability in community composition. Habitat (vernal pool, upland or riparian forest floor) and site of collection explained > 40%, but could be attributed to either niche or neutral processes. Hence, although niche dynamics may regulate fungal communities at the habitat scale, our evidence points towards neutral assembly of saprotrophic fungi on individual leaves, with energy availability constraining the taxa–area relationship.     

Reconciling niche and neutrality: the continuum hypothesis

In this study, we ask if instead of being fundamentally opposed, niche and neutral theories could simply be located at the extremes of a continuum. First, we present a model of recruitment probabilities that combines both niche and neutral processes. From this model, we predict and test whether the relative importance of niche vs. neutral processes in controlling community dynamics will vary depending on community species richness, niche overlap and dispersal capabilities of species (both local and long distance). Results demonstrate that niche and neutrality form ends of a continuum from competitive to stochastic exclusion. In the absence of immigration, competitive exclusion tends to create a regular spacing of niches. However, immigration prevents the establishment of a limiting similarity. The equilibrium community consists of a set of complementary and redundant species, with their abundance determined, respectively, by the distribution of environmental conditions and the amount of immigration.     

Grazing increases functional richness but not functional divergence in Tibetan alpine meadow plant communities

Plant community diversity and ecosystem function are conditioned by competition among co-occurring species for multiple resources. Previous studies suggest that removal of standing biomass by grazing decreases competition for light, but coincident grazing effects on competition for soil nutrients remain largely unknown in Tibetan rangelands where grazing tends to deplete soil phosphorus availability. We measured five functional traits indicative of plant productivity and stoichiometry leaf carbon concentration (LCC), leaf nitrogen concentration (LNC), leaf phosphorus concentration (LPC), specific leaf area (SLA), leaf dry matter content (LDMC) for component species of plant communities in grazed and ungrazed plots in five Tibetan alpine meadows. We examined the diversity of traits singly Rao index of functional diversity (FDrao) and in aggregate functional richness (FRic), functional divergence (FDiv), and functional evenness (FEve) in response to grazing. We tested whether foliar trait diversity increases with nutrient competition but decreases with light competition when competitive exclusion is reduced by grazing. The FDrao of LPC significantly increased under grazing, but FDrao for LCC, LNC and SLA tended to decrease. The FDrao of LDMC increased at the drier site but decreased at the wettest site. There was a strong negative association between increase in FDrao of LPC and decrease in soil nutrients, especially soil phosphorus availability. The FRic for all five traits together increased with species diversity following grazing, but neither FDiv nor FEve differed significantly between grazed and ungrazed plots at most sites. Grazing in Tibetan alpine meadows tends to increase competition for soil phosphorus while decreasing competition for light, resulting in an increase in the functional richness in grazed plant communities without any significant changes in the overall functional diversity of foliar traits. Our study highlights the potential importance of grazing mediated competition for multiple resources in alpine meadow ecosystems.     

The effects of long-term nitrogen loading on grassland insect communities

Just as long-term nitrogen loading of grasslands decreases plant species richness and increases plant biomass, we have found that nitrogen loading decreases insect species richness and increases insect abundances. We sampled 54 plots that had been maintained at various rates of nitrogen addition for 14 years. Total insect species richness and effective insect diversity, as well as herbivore and predator species richness, were significantly, negatively related to the rate of nitrogen addition. However, there was variation in trophic responses to nitrogen. Detritivore species richness increased as nitrogen addition increased, and parasitoids showed no response. Insect abundances, measured as the number of insects and insect biovolume (an estimate of biomass), were significantly, positively related to the rate of nitrogen addition, as were the abundances of herbivores and detritivores. Parasitoid abundance was negatively related to the rate of nitrogen addition. Changes in the insect community were correlated with changes in the plant community. As rates of nitrogen addition increased, plant species richness decreased, plant productivity and plant tissue nitrogen increased, and plant composition shifted from C₄ to C₃ grass species. Along this gradient, total insect species richness and effective insect diversity were most strongly, positively correlated with plant species richness. Insect biovolume was negatively correlated with plant species richness. Responses of individual herbivores varied along the nitrogen gradient, but numbers of 13 of the 18 most abundant herbivores were positively correlated with their host plant biomass. Although insect communities did not respond as strongly as plant communities, insect species richness, abundance, and composition were impacted by nitrogen addition. This study demonstrates that long-term nitrogen loading affects the entire food chain, simplifying both plant and insect communities. Received: 18 May 1999 / Accepted: 5 January 2000     

Temporal shifts and niche overlapping in <Emphasis Type="Italic">Copestylum</Emphasis> (Diptera,Syrphidae) communities reared in cactus species in a central Mexican scrubland

Semiarid scrubland communities are highly dynamic in terms of their species composition, abundance, and functioning, given the drastic changes in climate among seasons. Spatiotemporal patterns of saprophagous Copestylum (Diptera: Syrphidae) communities in different cactus species richness have not yet been studied, although seasonal changes and plant species richness have been shown to strongly impact the diversity and distribution of many insect communities in scrublands. We analyzed the impact of seasonality and of habitat type (disturbed and undisturbed) on Copestylum communities reared from cactus species at the Barranca de Metztitlán Biosphere Reserve, in central Mexico, by comparing their community structure between seasons and habitats, and assessing the contribution of diversity components for the total diversity of this genus. We also measured patterns of temporal niche overlap among hoverfly species considering their breeding medium. Seasonal variation influenced Copestylum community composition most significantly. Species richness and abundance of Copestylum were higher in the rainy season. Additive partitioning of diversity showed that the main component for species richness is beta diversity between seasons. We detected high niche overlap during the dry season and low overlap during the rainy season. This study provides evidence of temporal shifts in xeric hoverfly communities and suggests that the Copestylum species partition resources over time.