Patchy Distributions of Competitors Affect the Growth of a Clonal Plant When the Competitor Density Is High

Environments are patchy in not only abiotic factors but also biotic ones. Many studies have examined effects of spatial heterogeneity in abiotic factors such as light, water and nutrients on the growth of clonal plants, but few have tested those in biotic factors. We conducted a greenhouse experiment to examine how patchy distributions of competitors affect the growth of a rhizomatous wetland plant Bolboschoenus planiculmis and whether such effects depend on the density of the competitors. We grew one ramet of B. planiculmis in the center of each of the experimental boxes without competitors (Schoenoplectus triqueter), with a homogeneous distribution of the competitors of low or high density, and with a patchy distribution of the competitors of low or high density. The presence of competitors markedly decreased the growth (biomass, number of ramets, number of tubers and rhizome length) of the B. planiculmis clones. When the density of the competitors was low, the growth of B. planiculmis did not differ significantly between the competitor patches and competitor-free patches. However, when the density of the competitors was high, the growth of B. planiculmis was significantly higher in the competitor-free patches than in the competitor patches. Therefore, B. planiculmis can respond to patchy distributions of competitors by placing more ramets in competition-free patches when the density of competitors is high, but cannot do so when the density of competitors is low.     

Effects of soil nutrient heterogeneity on intraspecific competition in the invasive, clonal plant Alternanthera philoxeroides

Background and Aims

Fine-scale, spatial heterogeneity in soil nutrient availability can increase the growth of individual plants, the productivity of plant communities and interspecific competition. If this is due to the ability of plants to concentrate their roots where nutrient levels are high, then nutrient heterogeneity should have little effect on intraspecific competition, especially when there are no genotypic differences between individuals in root plasticity. We tested this hypothesis in a widespread, clonal species in which individual plants are known to respond to nutrient heterogeneity.

Methods

Plants derived from a single clone of Alternanthera philoxeroides were grown in the greenhouse at low or high density (four or 16 plants per 27·5 × 27·5-cm container) with homogeneous or heterogeneous availability of soil nutrients, keeping total nutrient availability per container constant. After 9 weeks, measurements of size, dry mass and morphology were taken.

Key Results

Plants grew more in the heterogeneous than in the homogeneous treatment, showing that heterogeneity promoted performance; they grew less in the high- than in the low-density treatment, showing that plants competed. There was no interactive effect of nutrient heterogeneity and plant density, supporting the hypothesis that heterogeneity does not affect intraspecific competition in the absence of genotypic differences in plasticity. Treatments did not affect morphological characteristics such as specific leaf area or root/shoot ratio.

Conclusions

Results indicate that fine-scale, spatial heterogeneity in the availability of soil nutrients does not increase competition when plants are genetically identical, consistent with the suggestion that effects of heterogeneity on competition depend upon differences in plasticity between individuals. Heterogeneity is only likely to increase the spread of monoclonal, invasive populations such as that of A. philoxeroides in China.     

水分异质性影响两种根茎型克隆植物赖草和假苇拂子茅的水分存储能力

水分在自然系统中呈异质性分布。有关水分异质性对克隆植物生长、形态和生理影响的研究已有大量的工作, 但是水分异质性对克隆植物存储能力, 尤其是水分存储能力影响的研究却十分缺乏。该文将两种根茎型克隆植物赖草(Leymus secalinus)和假苇拂子茅(Calamagrostis pseudophragmites)进行水分异质性和同质性实验处理, 探讨水分异质性对克隆植物水分存储能力、生长和形态的影响。在异质性水分处理下, 两种克隆植物的间隔子、枝和根的含水量均显著增加。两种克隆植物对水分异质性分布的适应策略有所不同, 赖草通过降低单个克隆分株的生长、提高芽的数量以应对水分异质性, 而假苇拂子茅通过增强整个分株种群的地下部分(根状茎、根和芽)生长来应对水分资源的异质性分布。水分储存能力的增强可以提高克隆植物适应水分异质性的能力。     

Soil Particle Heterogeneity Affects the Growth of a Rhizomatous Wetland Plant

Soil is commonly composed of particles of different sizes, and soil particle size may greatly affect the growth of plants because it affects soil physical and chemical properties. However, no study has tested the effects of soil particle heterogeneity on the growth of clonal plants. We conducted a greenhouse experiment in which individual ramets of the wetland plant Bolboschoenus planiculmis were grown in three homogeneous soil treatments with uniformly sized quartz particles (small: 0.75 mm, medium: 1.5 mm, or large: 3 mm), one homogeneous treatment with an even mixture of large and medium particles, and two heterogeneous treatments consisting of 16 or 4 patches of large and medium particles. Biomass, ramet number, rhizome length and spacer length were significantly greater in the treatment with only medium particles than in the one with only large particles. Biomass, ramet number, rhizome length and tuber number in the patchy treatments were greater in patches of medium than of large particles; this difference was more pronounced when patches were small than when they were large. Soil particle size and soil particle heterogeneity can greatly affect the growth of clonal plants. Thus, studies to test the effects of soil heterogeneity on clonal plants should distinguish the effects of nutrient heterogeneity from those of particle heterogeneity.     

Growth and competition in clonal plants-persistence of shoot populations and species diversity

This paper reviews studies on growth and size-structure dynamics of shoots and clones in clonal plants in comparison with those in non-clonal plants, and discusses the characteristics of clonal plants. The mode of competition between individuals (symmetric versus asymmetric, degree of competitive asymmetry), growth dynamics of individuals, allocation pattern between organs and spatial pattern of individuals are closely correlated with each other in non-clonal plant populations. Theoretical and field studies based on the diffusion model revealed that plants of “height-growth” type (mostly early-successional tree species) and plants of “diameter-growth” type (mostly late-successional tree species) tend to exhibit asymmetric competition and symmetric competition respectively. Moreover, asymmetrically competing plants show smaller effects of variation in individual growth rate and spatial pattern on the size-structure dynamics of the population than symmetrically competing plants. Thefefore, the spatial pattern of inviduals should be considered especially for plants undergoing symmetric competition. These results for non-clonal plants should have a significant implication also for the growth dynamics and competition in clonal plants. The mean growth rate of shoots [G(t,x) function] and hence the mode of competition between shoots differs among clonal plant species as in non-clonal plants. However, a large magnitude and size-independence (or slightly negative size-dependence) of the variation in growth rate of shoots [D(t,x) function], especially at the early stage in a growing season is a common characteristic of many clonal plant species, in contrast to the positively size-dependent variation in individual growth rate in non-clonal plants. This type of variation in shoot growth rate leads to the persistence of stable shoot populations even when the mean growth rate function is changed, and also in cases where the shoot population structure would be unstable in the absence of variation in growth rate. It is suggested that competition between clones is symmetric in most clonal plant species, which brings about small-scale spatio-temporal changes in species abundance and hence species diversity.     

Effects of Heterogeneous Competitor Distribution and Ramet Aggregation on the Growth and Size Structure of a Clonal Plant

Spatially heterogeneous distribution of interspecific competitors and intraspecific aggregation of offspring ramets may affect the growth and size structure of clonal plant populations, but these have been rarely studied. We conducted a greenhouse experiment in which we grew a population of eight offspring ramets (plants) of the stoloniferous clonal plant Hydrocotyle vulgaris aggregately or segregately in two homogeneous treatments with or without a competing grass Festuca elata and a heterogeneous treatment with a patchy distribution of the grass. In patchy grass treatments, H. vulgaris produced markedly more biomass, ramets and stolons in open patches (without grasses) than in grass patches, but displayed lower size variations as measured by coefficient of variation of biomass, ramets and stolons among the eight plants. In open areas, H. vulgaris produced statistically the same amounts of biomass and even more stolons and showed higher size variations in patchy grass treatments than in open (no grass) treatments. In grass areas, H. vulgaris grew much worse and displayed higher size variations in patchy grass treatments than in full grass treatments. Ramet aggregation decreased the growth of H. vulgaris in open treatments and in both open and grass patches in patchy grass treatments, but had little effect in full grass treatments. Ramet aggregation had little effect on size variations. Therefore, heterogeneous distribution of competitors can affect the growth and size structure of clonal plant populations, and ramet aggregation may decrease population growth when they grow in open environments or heterogeneous environments with a patchy distribution of interspecific competitors.     

Physiological integration affects growth form and competitive ability in clonal plants

Clonal plants translocate resources through spacers between ramets. Translocation can be advantageous if a plant occurs in heterogeneous environments (division of labour); however, because plants interact locally, any spatial arrangement of ramets generates some heterogeneity in light and nutrients even if there is no external heterogeneity. Thus the capacity of a clonal plant to exploit heterogeneous environment must operate in an environment where heterogeneity is partly shaped by the plant growth itself. Since most experiments use only simple systems of two connected ramets, plant-level effects of translocation are unknown. A spatially explicit simulation model of clonal plant growth, competition and translocation is used to identify whether different patterns of translocation have the potential to affect the growth form of the plant and its competitive ability. The results show that different arrangements of translocation sinks over the spacer system can completely alter clonal morphology. Both runners and clumpers can be generated using the same architectural rules by changing translocation only. The effect of translocation strongly interacts with the architectural rules of the plant growth: plants with ramets staying alive when a spacer is formed are much less sensitive to change in translocation than plants with ramets only at the tip. If translocation cost is low, translocating plants are in most cases better competitors than plants that do not translocate; the difference becomes stronger in more productive environments. Key traits that confer competitive ability are total number of ramet, and their fine-scale aggregation.Co-ordinating editor: J. Tuomi     

Morphological response to competition for light in the clonal Trifolium repens (Fabaceae)

? Premise of the study: Plant communities in temperate zones are dominated by clonal plants that can plastically modify their growth characteristics in response to competition. Given that plants compete with one another, and the implications this has for species coexistence, we conducted a study to assess how clonal species morphologically respond to competition for light depending on its intensity and heterogeneity, which are determined by the competitor species. ? Methods: We assessed the morphological response to competition for light of the clonal species Trifolium repens L. by measuring its growth performance, and vertical and horizontal growth traits. We used five competitive environments, i.e., one without competitor and four differing by their competitor species creating different conditions of competition intensity and heterogeneity. ? Key results: The morphological response of Trifolium repens to competition for light depended on the competitor identity. Competition intensity and heterogeneity, determined by competitor identity, had an interactive effect on most traits. The increase in petiole elongation and specific leaf area due to increased competition intensity was observed only at low to intermediate competition heterogeneity. Competition heterogeneity promoted the elongation of clone connections allowing space exploration. ? Conclusions: Our results demonstrated that the intensity and heterogeneity of competition, which depended on competitor identity, are of primary importance in determining the plastic response of Trifolium repens. This emphasizes that it is important to consider the fine-scale spatial distribution of individuals when studying their interactions within plant communities.     

Soil phosphorus heterogeneity and mycorrhizal symbiosis regulate plant intra-specific competition and size distribution

We investigated the interactive effects of soil phosphorus (P) heterogeneity, plant density and mycorrhizal symbiosis on plant growth and size variability of Trifolium subterraneum. We set up mesocosms (trays 49Ꮉ cm and 12 cm deep) with the same amount of available P, but distributed either homogeneously or heterogeneously, in randomly arranged cells (7ǻ cm each) with high or low available P. The trays were planted with either 1 or 4 seedlings of T. subterraneum per cell. Half of the trays were inoculated with spores of the mycorrhizal fungus Gigaspora margarita. We harvested the plants when leaves just started to overlap, 8 weeks after planting. Plants growing in high P cells had the lowest percentage infection, but the highest mean shoot and root biomass and root length. The mean size of the plants in each cell was determined mainly by local P concentration. However, in plants growing in high density, low P cells, ca. 20% of the variability in plant biomass was explained by the number of adjacent cells with high P. Patchy trays had the highest total shoot biomass, independently of mycorrhizal infection or plant density. Inoculated trays (M) had higher total shoot biomass and relative competition intensity (measured as reduction in plant biomass due to increased density) than non-inoculated trays (NM). Plant density reduced the plant response to mycorrhizal infection, and its effect was independent of P distribution. All populations growing in patchy trays, and low density mycorrhizal ones, had the highest plant-size inequality, presumably because patchy distribution of P and mycorrhizal infection increased competitive asymmetry. We conclude that mycorrhizal symbiosis has the potential to strongly influence plant population structure when soil nutrient distribution is heterogeneous because it promotes pre-emption of limiting resources.     

Spatial covariance in resources affects photosynthetic rate and water potential,but not the growth of Glechoma longituba fragments

Glechom longituba is a widespread clonal plant and usually experiences diverse patchiness of its growing sites. The hypothesis was tested that spatial covariance in resources differentially affects the growth and physiology of G. longituba. Plants were exposed to two patchy habitats where above- and below-ground resources were either positively or negatively associated, and to four control habitats. When equal amounts of resources were given, G. longituba fragments from two patchy habitats had similar biomass, root weight ratio, leaf weight ratio, fluorescence yield, specific petiole length, and specific root length; fragments under reciprocal patchiness significantly increased photosynthetic rate and decreased leaf water potential. Biomass of clones grown in patchy habitats was equal or less than that of counterparts from the control habitats, not supporting the notion that clonal growth is more advantageous in patchy habitats than in uniform habitats. In addition, no evidence was detected for spatial division of labour because biomass allocation to roots and leaves was similar in patchy habitats compared with the control habitats.     

A negative heterogeneity–diversity relationship found in experimental grassland communities

Recent meta-analyses and simulation studies have suggested that the relationship between soil resource heterogeneity and plant diversity (heterogeneity–diversity relationship; HDR) may be negative when heterogeneity occurs at small spatial scales. To explore different mechanisms that can explain a negative HDR, we conducted a mesocosm experiment combining a gradient of soil nutrient availability (low, medium, high) and scale of heterogeneity (homogeneous, large-scale heterogeneous, small-scale heterogeneous). The two heterogeneous treatments were created using chessboard combinations of low and high fertility patches, and had the same overall fertility as the homogeneous medium treatment. Soil patches were designed to be relatively larger (156 cm²) and smaller (39 cm²) than plant root extent. We found plant diversity was significantly lower in the small-scale heterogeneous treatment compared to the homogeneous treatment of the same fertility. Additionally, low fertility patches in the small-scale heterogeneous treatment had lower diversity than patches of the same size in the low fertility treatment. Shoot and root biomass were larger in the small-scale heterogeneous treatment than in the homogeneous treatment of the same fertility. Further, we found that soil resource heterogeneity may reduce diversity indirectly by increasing shoot biomass, thereby enhancing asymmetric competition for light resources. When soil resource heterogeneity occurs at small spatial scales it can lower plant diversity by increasing asymmetric competition belowground, since plants with large root systems can forage among patches and exploit soil resources. Additionally, small-scale soil heterogeneity may lower diversity indirectly, through increasing light competition, when nutrient uptake by competitive species increases shoot biomass production.     

Spatial heterogeneity in light supply affects intraspecific competition of a stoloniferous clonal plant

Spatial heterogeneity in light supply is common in nature. Many studies have examined the effects of heterogeneous light supply on growth, morphology, physiology and biomass allocation of clonal plants, but few have tested those effects on intraspecific competition. In a greenhouse experiment, we grew one (no competition) or nine ramets (with intraspecific competition) of a stoloniferous clonal plant, Duchesnea indica, in three homogeneous light conditions (high, medium and low light intensity) and two heterogeneous ones differing in patch size (large and small patch treatments). The total light in the two heterogeneous treatments was the same as that in the homogeneous medium light treatment. Both decreasing light intensity and intraspecific competition significantly decreased the growth (biomass, number of ramets and total stolon length) of D. indica. As compared with the homogeneous medium light treatment, the large patch treatment significantly increased the growth of D. indica without intraspecific competition. However, the growth of D. indica with competition did not differ among the homogeneous medium light, the large and the small patch treatments. Consequently, light heterogeneity significantly increased intraspecific competition intensity, as measured by the decreased log response ratio. These results suggest that spatial heterogeneity in light supply can alter intraspecific interactions of clonal plants.     

Interspecific competition changes reproductive output but does not increase reproductive costs in a grassland perennial

In plants, it is hypothesized that allocation trade-offs may appear only when expenditures like seed production are high or external resources are scarce. In this study, we tested whether reproductive costs are more pronounced under enhanced interspecific competition.In a common garden, we investigated phenotypic correlations between sexual reproduction, clonal growth and storage structures in the grassland perennial, Succisa pratensis. During the past 50 years, habitats of this species have faced an expansion of clonal grasses that increase competition intensity. We simulated this process by growing five populations of Succisa from high- and low-production habitats with its clipped and non-clipped competitor, Agrostis capillaris. In addition, we experimentally removed flower heads of Succisa plants from one population grown with and without A. capillaris.We demonstrated costs of sexual reproduction by flower-head removal (resulting in increased plant size and relative allocation to belowground structures) but not by phenotypic correlations. We found no evidence that reproductive costs increase in a competitive environment and the opposite pattern was shown in both approaches used. However, high competition intensity reduced relative investment to flower-head production. In plants from low-production habitats, competition also reduced the absolute number of flower heads and belowground biomass as a result of smaller plant size. We assume that populations from low-production habitats are more prone to extinction as they have a reduced likelihood of local persistence and of escape to more suitable habitats during advancing succession.     

Effect of local spatial plant distribution and conspecific density on bumble bee foraging behaviour

1. Size variations in pollinator populations may modify competitive interactions among foragers. Competition among pollinators has been shown to lead to one of two contrasting behaviours: either specialisation to the most profitable plant species or generalisation to several species. When foraging, pollinators are also confronted with heterogeneity in the spatial distribution of plant resources. Because variations in both the forager density and plant spatial distribution can affect pollinator behaviour, focus was on the interactive effect of these two factors. 2. Bumble bee (Bombus terrestris L.) individuals were trained on a focal species (Lotus corniculatus L.) and experimentally tested whether variations in the forager density (two or six bumble bees foraging together), plant community spatial distribution (two plant species: L. corniculatus and Medicago sativa, which were either patchily or randomly distributed), and their interaction modified bumble bee foraging behaviour. 3. It was shown that when confronted with a high forager density, bumble bees focused their visits towards the most familiar species, especially when foraging under a random plant distribution. These modifications affected the fruiting of the focal plant species, with a significantly lower reproductive success under low density/patchy conditions. 4. This study demonstrates that the foraging decisions of bumble bees are influenced by variations in both the conspecific density and plant spatial distribution. Given the increasing impact of human activities on plant‐pollinator communities, this raises the question of the potential implications of these results for plant communities in natural conditions when confronted with variations in the pollinator density and spatial distribution of plants.     

Clonal Growth in Plants in Relation to Resource Heterogeneity:Foraging Behavior

In nature, essential resources for organisms, such as food for animals and light, water and nutrients for plants, are usually heterogeneously distributed, even at very small scale. As a result, all organisms, particularly plants mostly sessile, have a difficulty in acquiring essential resources from their environments. Animals express various types of foraging behavior to capture heterogeneously distributed essential foods. Clonal growth ( a vegetative reproductive process where by more than one individual of identical genetic composition is formed ) provides clonal plant not only with many "mouths" at different spatial positions, but also with a large spacial movability. As a clonal plant grows in environments characterized by a small-scale resource heterogeneity, its inter ramet connection permits a resource-sharing among the connected tamers. In addition, it may also allow certain ramets to respond locally and non-locally to resousce heterogeneity. This may lead to a division of labor among the connected ramets and a selective placement of ramets in favorable micro-habitats. Together these may enhance exploitation of resource heterogeneity by clonal plants, and in turn greatly contribute to maintenance or improvement of fitness. Such a behavior of clonal plants, expressed in heterogeneous environments, is to a large extent comparable to that of animals. Therefore, it has been considered as foraging behavior in clonal plants. More recently, it has been observed that phenotypic plasticity of clonal plants, which is relevant to foraging behavior, varies among species, types of genet architecture as well as among types of plants habitats. Foraging in clonal plants and its diversity have been receiving increasingly intensive investigations.     

Plant size and spatial pattern in a natural population of Myosotis micrantha

Abstract. We examined spatial distributions and plant sizes along a transect through a natural population of a winter annual, Myosotis micrantha. A size hierarchy existed, as indicated by high values of Gini coefficients of inequality for plant mass and correlated measures. Plants with no immediate conspecific neighbors were larger than plants with one or more near neighbors, suggesting that competition from near neighbors depressed plant size. However, there was strong positive spatial autocorrelation in plant size: large plants were associated with large neighbors and small ones with small neighbors. Plant size was also positively correlated with the combined biomass of near neighbors. The population formed a two-phase mosaic of patches of relatively large plants alternating with patches of smaller plants. The data suggest that individual plants compete with conspecifics, but the effects of competition are symmetrical. The most likely explanations for this spatially structured size hierarchy are variation in plant density, patchy distribution of resources, or a combination of the two.     

Effects of Spatial Scale of Soil Heterogeneity on the Growth of a Clonal Plant Producing Both Spreading and Clumping Ramets

Soil nutrients are commonly heterogeneously distributed at different spatial scales. Although numerous studies have tested the effects of soil nutrient heterogeneity on growth of clonal plants producing either spreading ramets or clumping ramets, no study has examined the effects on the growth of clonal plants producing both spreading and clumping ramets and how spatial scale affects such effects. To test these effects, clones of Buchloe dactyloides, a stoloniferous clonal plant that produces both clumping and spreading ramets, were grown in six heterogeneous environments with different patch sizes and one homogeneous environment containing the same quantity of nutrients. Total biomass, total number of ramets, number of clumping ramets, number of spreading ramets, spacer length, or root:shoot ratio of the whole plants did not differ significantly among the seven treatments. However, at the patch level there were significant effects of patch size by nutrient level on biomass, number of ramets, number of spreading ramets, and number of clumping ramets, and these four variables were significantly larger in the nutrient-rich patches than in the nutrient-poor patches in the heterogeneous treatment with the largest patch size, but not in the other five heterogeneous treatments with smaller patch sizes. Neither nutrient level nor patch size significantly affected spacer length or root:shoot ratio. Based on our results, we propose that B. dactyloides can efficiently exploit nutrient-rich patches by a plastic response of clumping ramets and spreading ramets at larger spatial scales of soil heterogeneity but not at smaller ones.     

Optimal root proliferation strategies: the roles of nutrient heterogeneity, competition and mycorrhizal networks

Background and Aims Plants proliferate roots in order to acquire nutrients, typically contending with heterogeneous resources and competing neighbours. A mathematical model was developed to identify optimal root proliferation strategies in patchy nutrient environments. The impact of joining mycorrhizal networks was also assessed. Methods A simple model of growth and competition in one spatial dimension was implemented within a genetic algorithm to obtain optimal proliferation strategies under different scenarios of resource distribution, and in the presence or absence of local competition and large-scale mycorrhizal networks. Results A strong proliferation response emerged for isolated plants in heterogeneous environments with low resources, and also for plants growing in competition. Even in statistically homogeneous environments, the presence of competition conferred a selective advantage to plants proliferating in the direction of the most recently acquired patch. In the presence of mycorrhizal networks, the optimal strategy switched from symbiosis to proliferation driven growth as the relative cost of acquiring resources through the networks increased. Conclusions The optimal proliferation response in a given scenario was governed by a hierarchy of factors: resource levels and distribution; the presence or absence of competition; and the marginal benefit of obtaining resources via symbiotic relationships with mycorrhizas.     

On the ecological and evolutionary significance of storage in clonal plants

Environmental heterogeneity has received wide attention in clonal plant research over the last decade. Most studies have focussed on the effects of spatial variation in environmental conditions on the performance of ramets and genets, while the effects of temporal heterogeneity have only occasionally been studied. As a consequence, our current knowledge about functional responses of clonal plants to habitat patchiness is biased towards spatial aspects of environmental heterogeneity. Nevertheless, temporal changes in biotic and abiotic conditions do occur in most natural habitats, and they are very likely to affect plant growth and performance, and to create positive selection pressures on traits that can buffer plants against unfavorable consequences of this variability. Storage of resources is a widespread phenomenon in clonal plant species. Typical clonal structures such as stolons, rhizomes and hibernacles serve as storage organs. However, the ecologic significance of storage in clonal plant structures remains partly unclear. We suggest that the lack of understanding with respect to resource storage in clonal plants be related to our poor knowledge of ecologic implications of temporal habitat heterogeneity in natural environments. Resource storage can be understood as a safety measure against temporal changes in the growing conditions of plants. This paper summarizes existing information about the ecologic relevance of storage in clonal plants and it tries to develop a framework for further investigation of resource storage as a strategy to enhance the performance of clonal plants in temporally variable environments.     

Resource heterogeneity, soil fertility, and species diversity: effects of clonal species on plant communities

Spatial heterogeneity in soil resources is widely thought to promote plant species coexistence, and this mechanism figures prominently in resource-ratio models of competition. However, most experimental studies have found that nutrient enhancements depress diversity regardless of whether nutrients are uniformly or heterogeneously applied. This mismatch between theory and empirical pattern is potentially due to an interaction between plant size and the scale of resource heterogeneity. Clonal plants that spread vegetatively via rhizomes or stolons can grow large and may integrate across resource patches, thus reducing the positive effect of small-scale resource heterogeneity on plant species richness. Many rhizomatous clonal species respond strongly to increased soil fertility, and they have been hypothesized to drive the descending arm of the hump-shaped productivity-diversity relationship in grasslands. We tested whether clonals reduce species richness in a grassland community by manipulating nutrient heterogeneity, soil fertility, and the presence of rhizomatous clonal species in a 6-year field experiment. We found strong and consistent negative effects of clonals on species richness. These effects were greatest at high fertility and when soil resources were applied at a scale at which rhizomatous clonals could integrate across resource patches. Thus, we find support for the hypothesis that plant size and resource heterogeneity interact to determine species diversity.