Large herbivores maintain a two‐phase herbaceous vegetation mosaic in a semi‐arid savanna

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Soil patchiness in juniper-sagebrush-grass communities of central Oregon

This study compared the sizes, spacings and properties (soil moisture, pH, nitrogen, soil arthropods and VAM) of soil resource islands and bare patches in sagebrush-grass communities invaded by western juniper versus those without juniper. We analyzed 1000 surface soil samples taken from nine 50-m radius circular plots sampled in December of 1991 and May of 1992 on ‘The Island’, one of the few undisturbed areas of sagebrush-grass shrubland in Oregon. Spatial structure was interpreted from correlograms (Moran's I) and standardized semivariograms. The presence of juniper was associated with increased bare area and smaller, more widely spaced grass and sagebrush plants. Soil arthropod numbers and biomass in plots with juniper were only roughly one-fifth of those in sagebrush-grass plots in December. The dominant soil pattern in both sagebrush-grass and juniper-sagebrush-grass plots was regularly-distributed patches spanning a range of sizes and spacings. Plots with juniper had greater patchiness at shorter lags (<3 m), and patchiness was more developed for soil moisture, net nitrification, and net N mineralization, whereas sagebrush-grass plots had greater patchiness at longer lags (3 – 9 m) and patchiness was more developed for NO₃–N, arthropod numbers and biomass. These differences in soil patterns with and without juniper indicate that juniper responds to, or causes, changes in the size of resource islands under sage and grass when it invades sage-grass communities. This revised version was published online in June 2006 with corrections to the Cover Date.     

太行山南麓山区不同植被恢复类型土壤理化和细根结构特征

在干旱半干旱地区,由于水分匮乏、土壤贫瘠等因素,将形成一定的裸地斑块,而这些斑块极易造成水土流失、滑坡等灾害,而具有不同植被覆盖的林地则能有效的保持水土。为完善干旱半干旱地区不同植被恢复类型下土壤理化和细根特征,选择太行山南麓山区具有代表性的裸露地、草地、荆条地、侧柏地、栓皮地和刺槐地等植被恢复类型,比较了各植被恢复类型下的土壤养分、粒径及细根状况等差异。研究表明:1)相对于裸露地,有植被覆盖的植被恢复类型拥有良好的土壤及细根状况。2)在不同植被类型中,刺槐林的有效氮转化速率较高;侧柏林有较高的细根参数;草地能够提高土壤中可吸收的磷组分。3)林地类型和土层均对土壤中含水率、黏粒、细根生物量和比根长产生极显著影响(P0.001)。4)各植被类型的对于土壤斑块的利用能力不同;不同植被类型中土壤及细根状况变化量具有一定的相似性,研究为生态恢复中植被类型的合理布局提供了新思路。     

Sex‐related spatial patterns of Poa ligularis in relation to shrub patch occurrence in northern Patagonia

Abstract. Poa ligularis is a dioecious species and a valuable forage plant which is widespread in the arid steppe of northern Patagonia (Argentina). The vegetation in these areas consists of a system of perennial plant patches alternating with bare soil areas defining contrasting micro‐environments. We hypothesized that (1) male and female individuals of P. ligularis are spatially segregated in different micro‐environments, (2) the intensity of spatial segregation of sexes depends on plant structure and (3) spatial segregation of sexes is enhanced by competitive interactions between the sexes within the vegetation patches. We analysed the spatial distribution of female and male individuals in relation to the spatial pattern of vegetation in two areas differing in their vegetation structure. The location of P. ligularis within patches where either male, female or both sexes occurred was also analysed. The results indicate that different patterns of spatial distribution of sexes of P. ligularis may be found at the community level depending on the dominant life forms and geometric structure of plant patches. Where patches are of a lower height, with a high internal patch cover, individuals of both sexes are concentrated within patch canopies. In sites characterized by large, tall patches and less internal patch cover suitable microsites for female and male P. ligularis occur both within and outside the patch with males located at further distances from the patch edge. Where the patch is large and tall enough to allow the establishment of males and females at relatively high numbers, males occupy the patch periphery or even colonize the interpatch bare soil. These spatial patterns are consistent with selective traits in which females better tolerate intraspecific competition than males, while males tolerate wider fluctuations in the physical environment (soil moisture, nitrogen availability, wind intensity, etc.).     

Spatial heterogeneity and irreversible vegetation change in semiarid grazing systems

Recent theoretical studies have shown that spatial redistribution of surface water may explain the occurrence of patterns of alternating vegetated and degraded patches in semiarid grasslands. These results implied, however, that spatial redistribution processes cannot explain the collapse of production on coarser scales observed in these systems. We present a spatially explicit vegetation model to investigate possible mechanisms explaining irreversible vegetation collapse on coarse spatial scales. The model results indicate that the dynamics of vegetation on coarse scales are determined by the interaction of two spatial feedback processes. Loss of plant cover in a certain area results in increased availability of water in remaining vegetated patches through run-on of surface water, promoting within-patch plant production. Hence, spatial redistribution of surface water creates negative feedback between reduced plant cover and increased plant growth in remaining vegetation. Reduced plant cover, however, results in focusing of herbivore grazing in the remaining vegetation. Hence, redistribution of herbivores creates positive feedback between reduced plant cover and increased losses due to grazing in remaining vegetated patches, leading to collapse of the entire vegetation. This may explain irreversible vegetation shifts in semiarid grasslands on coarse spatial scales.     

Spatial heterogeneity in the herbaceous layer of a semi-arid savanna ecosystem

Despite increasing recognition of the role spatial pattern can play in ecosystem function, few studies have quantified spatial heterogeneity in savanna ecosystems. The spatial distribution of herbaceous biomass and species composition was measured across three scales in a semi-arid savanna in central Kenya, and patterns were related to environmental variables at different scales. Herbaceous biomass declined across a rainfall gradient and from upper to lower topographic positions, but variation within a site (across 5–50 m) was similar in magnitude to among-site variation associated with rainfall and topography. Geostatistical analyses showed that patchiness at scales of 5–25 m explained 20% of total variation in herbaceous biomass. This pattern arose from the presence of both 5–10-m diameter patches containing high herbaceous biomass (> 170 g m^–2) and 5–10-m diameter patches characterized by nearly bare soil surfaces (< 40 g m^–2). Patch structure was contingent on topography, with larger bare patches at ridgeline and upper hillslope positions. Grass species distributions showed the greatest degree of patch structure and species turnover across distances of 5–45 m. Additional community variation was associated with topography, with minimal variation in species composition across the rainfall gradient. Pattern diversity significantly exceeded levels reported for four other grassland ecosystems, suggesting fundamental differences in local processes generating spatial pattern. It is hypothesized that heterogeneously distributed grazing pressure, interacting with the distribution of shrub canopies, is an important factor generating such high levels of small-scale patch structure in this savanna.     

Root proliferation strategies and exploration of soil patchiness in arid communities

Soil patchiness is a key feature of arid rangelands. As root proliferation contributes to soil exploration and resource uptake, it is ecologically relevant to understand how species respond to soil heterogeneity and coexist. Campbell et al.'s influential 1991 hypothesis proposes that dominant species deploy root systems (scale) that maximize soil volume explored. Instead, subordinate species show accurate root systems that exclusively proliferate in nutrient‐rich patches (precision). After many experiments under controlled conditions, the generality of this hypothesis has been questioned but a field perspective is necessary to increase realism in the conceptual framework. We worked with a guild of perennial graminoid species inside a grazing exclosure in an arid Patagonian steppe, a model system for ecological studies in arid rangelands for four decades. We buried root traps in bare ground patches with sieved soil, with or without a pulse of nitrogen addition, to measure specific root biomass and precision at 6 and 18 months after burial. We also estimated scale (root density) in naturally established plants, and root decomposition in litter bags. Several species grew in root traps. Dominant species showed the highest root biomass (in both harvests) and scale. Subordinate species grew more frequently with nitrogen addition and showed lower biomass and scale. Similar total root biomass was found with and without nitrogen addition. Species differed in root decomposition, but correcting species biomass by decomposition did not change our conclusions. We did not find a relation between scale and precision, indicating that Campbell's hypothesis is probably not supported in this Patagonian steppe. Soil resource acquisition differences probably do not utterly explain the coexistence of dominant and subordinate species because the steppe is also affected by large herbivore grazing. We propose that root proliferation in this steppe is the result of the interaction between individual density in the community and specific root growth rates.     

Spatial patterns of ground vegetation, soil microbial biomass and activity in a mixed spruce-birch stand

Trees directly and indirectly influence the above- and below-ground environment, and can be expected to modify the spatial patterns of organisms associated with the forest floor. This study aimed to examine the effects of a coniferous (Picea abies) and a broad-leaved (Betula pubescens) tree species on the spatial pattern of ground vegetation and soil microbial properties in a mixed stand in central Sweden. I have characterised the species composition of ground vegetation, soil microbial biomass and activity, photosynthetic active radiation (PAR), soil water content and soil pH in the stand, and tested whether the spatial patterns of these variables were related to the positioning of trees. Geostatistics were used to describe the spatial variation in ground vegetation, soil mirobiological properties and the soil surface environment. PAR, soil water content and the cover of the moss Brachytecium reflexum and associated herb species decreased with the influence of spruce trees. Microbial biomass, measured as the amount of phospholipid fatty acids, decreased with spruce influence but increased with the influence of birch trees. Microbial respiration was not affected by spruce but increased with the influence of birch. Ground vegetation and microbial respiration, which were influenced by one tree species only, aggregate on a scale of 4-5 m, corresponding fairly well with patches of a single tree species. Soil microbial biomass, which was affected by both tree species, aggregated on a scale of 7-8 m. roughly corresponding to the distance between patches of spruce and birch trees respectively. I suggest that spruce trees influenced vegetation mainly through shading, and that a difference in the availability of organic matter under birch and spruce trees caused spatial variation in microbial biomass and activity. Thus, spatial patterns in ground vegetation and soil microbial properties may develop in a mixed forest of coniferous-broad leaved trees, as a result of the difference in influence of tree species and nested variation associated with the arrangement of the trees.     

宁夏东部荒漠草原向灌丛地人为转变过程中土壤胞外酶活性响应

以宁夏东部荒漠草原-灌丛地典型镶嵌体内部荒漠草地、草地边缘、灌丛边缘、灌丛地为对象,对各样地植丛和空斑下土壤特性及6种土壤胞外酶活性(纤维二糖水解酶、β-1,4-木糖苷酶、β-1,4葡萄糖苷酶、β-1,4-乙酰基氨基葡萄糖苷酶、亮氨酸氨基肽酶和碱性磷酸酶)进行分析,研究荒漠草原向灌丛地人为转变过程中胞外酶的响应特征。结果表明: 荒漠草原向灌丛地转变过程中,土壤水分、有机碳、全氮、全磷、微生物生物量碳、微生物生物量氮均显著降低,且灌丛地显著低于草地26.0%～88.5%;除草地边缘土壤水分、有机碳空斑略高于植丛外,其他指标均表现为各样地植丛显著高于空斑3.9%～82.3%。6类土壤胞外酶活性在转变过程中均呈下降趋势,降幅为22.1%～82.4%,其中亮氨酸氨基肽酶和碱性磷酸酶降低最为显著,分别降低82.4%和75.5%;除灌丛地β-1,4-乙酰基氨基葡萄糖苷酶在空斑显著高于植丛外,其他胞外酶活性均表现为各样地植丛高于空斑10.7%～42.7%;转变过程中6类胞外酶活性之间呈显著正相关,且均与土壤特性呈不同程度正相关,其中各类土壤胞外酶活性对土壤微生物生物量碳、氮及全氮响应较为积极。     

Relationship between small-scale structural variability and simulated vegetation productivity across a regional moisture gradient in southern Africa

The observed variability in vegetation structure within landscapes was used as the basis for model estimates of the range of annual productivity of landscape patches at four sites along a moisture gradient in southern Africa ranging from 879 to 365 mm mean annual rainfall. Principal components of patch‐scale variability in leaf area, woody biomass and vertical leaf profiles were derived from intensive characterization of the small‐scale spatial structure of woody vegetation at each site. For each site, the mean and extremes of the principal component distribution parameterized an ecophysiology model of vegetation productivity. Vegetation was most heterogeneous at intermediate locations along the rainfall gradient. Variability in vegetation structure led to a range of annual productivity within one site (600 mm) that accounted for 68% of the total range in mean productivity across all sites. Patch‐scale estimates of tree productivity were found to be primarily correlated to annual rainfall (r²=0.66, P=0.001) and not woody leaf area (r²=0.01, P=0.75), while grass productivity was found to be related to values of woody leaf area (r²=0.77, P<0.001) and not annual rainfall (r²=0.11, P=0.29). This result indicates that life‐form interactions have a significant role in controlling vegetation productivity across the rainfall gradient. The findings of this study emphasize the importance of considering heterogeneity rather than mean structure when modeling productivity, particularly when considering dynamic vegetation structure, where differences between landscape patches may not be well represented in the mean structure.     

Herbivores as architects of savannas: inducing and modifying spatial vegetation patterning

In this paper, we address the question whether and through which mechanisms herbivores can induce spatial patterning in savanna vegetation, and how the role of herbivory as a determinant of vegetation patterning changes with herbivore density and the pre-existing pattern of vegetation. We thereto developed a spatially explicit simulation model, including growth of grasses and trees, vertical zonation of browseable biomass, and spatially explicit foraging by grazers and browsers. We show that herbivores can induce vegetation patterning when two key assumptions are fulfilled. First, herbivores have to increase the attractiveness of a site while foraging so that they will revisit this site, e.g. through an increased availability or quality of forage. Second, foraging should be spatially explicit, e.g. when foraging at a site influences vegetation at larger spatial scales or when vegetation at larger spatial scales influences the selection and utilisation of a site. The interaction between these two assumptions proved to be crucial for herbivores to produce spatial vegetation patterns, but then only at low to intermediate herbivore densities. High herbivore densities result in homogenisation of vegetation. Furthermore, our model shows that the pre-existing spatial pattern in vegetation influences the process of vegetation patterning through herbivory. However, this influence decreases when the heterogeneity and dominant scale of the initial vegetation decreases. Hence, the level of adherence of the herbivores to forage in pre-existing patches increases when these pre-existing patches increase in size and when the level of vegetation heterogeneity increases. The findings presented in this paper, and critical experimentation of their ecological validity, will increase our understanding of vegetation patterning in savanna ecosystems, and the role of plant–herbivore interactions therein.     

Clonal Plasticity in Response to Reciprocal Patchiness of Light and Nutrients in the Stoloniferous Herb Glechoma longituba L.

Interconnected ramets of clonal plants can functionally specialize in the uptake of resources from aboveground and/or underground sources. Ramet pairs of the clonal stoloniferous herb Glechoma Iongltuba L. were grown In spatially heterogeneous environments with complementary availability of light and nutrients. Plasticity with respect to root-shoot ratio, fitness-related traits （biomass, number of ramets and dry weight per ramet）, morphological traits （lamina area, root length） were experimentally examined. The aim was to understand the adaptation of G. Iongltuba to an environment with reciprocal patchiness of light and soil nutrients by plasticity In Its root-shoot ratio and clonal morphology. The results showed that the performance of ramets growing In patches with high light Intensity and low soil nutrients into the adjacent opposite patches was Increased in terms of fitness-related traits. However, the performance of those from patches with low light Intensity and high soil nutrients into the adjacent opposite patches was not changed. The root-shoot ratio and clonal morphology were plastic. Morphological traits such as lamina area and root length were altered In a way that enhanced the capture of light resources and soil nutrients. Apparent reciprocal resource transport between the ramets In an environment of reciprocal patchiness of resources can enhance the growth of ramets with complementary resource deficiencies.     

Salt stress limitation of seedling recruitment in a salt marsh plant community

Summary Seedling recruitment in salt marsh plant communities is generally precluded in dense vegetation by competition from adults, but is also relatively rare in disturbance-generated bare space. We examined the constraints on seedling recruitment in New England salt marsh bare patches. Under typical bare patch conditions seed germination is severely limited by high substrate salinities. We examined the germination requirements of common high marsh plants and found that except for one notably patch-dependent fugitive species, the germination of high marsh plants is strongly inhibited by the high soil salinities routinely encountered in natural bare patches. Watering high marsh soil in the greenhouse to alleviate salt stress resulted in the emergence of up to 600 seedlings/225 cm². The vast majority of this seed bank consisted of Juncus gerardi, the only common high marsh plant with high seed set. We tested the hypothesis that salt stress limits seedling contributions to marsh patch secondary succession in the field. Watering bare patches with fresh water partially alleviated patch soil salinities and dramatically increased both the emergence and survival of seedlings. Our results show that seedling recruitment by high marsh perennial turfs is limited by high soil salinities and that consequently their population dynamics are determined primarily by clonal growth processes. In contrast, populations of patch-dependent fugitive marsh plants which cannot colonize vegetatively are likely governed by spatially and temporally unpredictable windows of low salinities in bare patches.     

Scale-dependent bi-trophic interactions in a semi-arid savanna: how herbivores eliminate benefits of nutrient patchiness to plants

The scale of resource heterogeneity may influence how resources are locally partitioned between co-existing large and small organisms such as trees and grasses in savannas. Scale-related plant responses may, in turn, influence herbivore use of the vegetation. To examine these scale-dependent bi-trophic interactions, we varied fertilizer [(nitrogen (N)/phosphorus (P)/potassium (K)] applications to patches to create different scales of nutrient patchiness (patch size 2 × 2 m, 10 × 10 m, or whole-plot 50 × 50 m) in a large field experiment in intact African savanna. Within-patch fertilizer concentration and the total fertilizer load per plot were independently varied. We found that fertilization increased the leaf N and P concentrations of trees and grasses, resulting in elevated utilization by browsers and grazers. Herbivory off-take was particularly considerable at higher nutrient concentrations. Scale-dependent effects were weak. The net effect of fertilization and herbivory was that plants in fertilized areas tended to grow less and develop smaller rather than larger standing biomass compared to plants growing in areas that remained unfertilized. When all of these effects were considered together at the community (plot) level, herbivory completely eliminated the positive effects of fertilization on the plant community. While this was true for all scales of fertilization, grasses tended to profit more from coarse-grained fertilization and trees from fine-grained fertilization. We conclude that in herbivore-dominated communities, such as the African savanna, nutrient patchiness results in the herbivore community profiting rather more than the plant community, irrespective of the scale of patchiness. At the community level, the allometric scaling theory’s prediction of plant—and probably also animal—production does not hold or may even be reversed as a result of complex bi-trophic interactions.     

Plant-Herbivore Interaction and Its Consequences for Succession in Wetland Ecosystems: A Modeling Approach

Herbivore grazing is increasingly used as a management tool to prevent the dominance of vegetation by tall grasses or trees. In this report, a model is described that is used to analyze plant-herbivore interactions and their scaling up to landscape scale. The model can be used to predict effects of herbivory on vegetation development. The model is an ecosystem model including modules for carbon and nitrogen cycling through plants, soil organic matter, and atmosphere. Plants compete for light and nitrogen. An herbivory module is included that implements selective foraging by a herbivore in a spatially heterogeneous area. Simulations were done to analyze the effects of herbivore density on vegetation dynamics, to analyze the impact of soil fertility on maximum herbivore density, and to analyze effects of herbivore density on landscapes. Two important points come forward from the model. Maximum herbivore abundance shows a hump-shaped curve along a soil fertility gradient. At higher soil fertility, light competition becomes more important. Herbivory interferes with plant competition, giving the tall, less palatable species a competitive advantage and thereby reducing the food quality and availability and hence the carrying capacity of the area. At a landscape scale, herbivory leads to increased heterogeneity. This increased heterogeneity may increase carrying capacity. The implications of these points for nature management are discussed. Received 13 May 1998; accepted 23 November 1998.     

Vertical distribution of the soil microbiota along a successional gradient in a glacier forefield

Spatial patterns of microbial communities have been extensively surveyed in well‐developed soils, but few studies investigated the vertical distribution of micro‐organisms in newly developed soils after glacier retreat. We used 454‐pyrosequencing to assess whether bacterial and fungal community structures differed between stages of soil development (SSD) characterized by an increasing vegetation cover from barren (vegetation cover: 0%/age: 10 years), sparsely vegetated (13%/60 years), transient (60%/80 years) to vegetated (95%/110 years) and depths (surface, 5 and 20 cm) along the Damma glacier forefield (Switzerland). The SSD significantly influenced the bacterial and fungal communities. Based on indicator species analyses, metabolically versatile bacteria (e.g. Geobacter) and psychrophilic yeasts (e.g. Mrakia) characterized the barren soils. Vegetated soils with higher C, N and root biomass consisted of bacteria able to degrade complex organic compounds (e.g. Candidatus Solibacter), lignocellulolytic Ascomycota (e.g. Geoglossum) and ectomycorrhizal Basidiomycota (e.g. Laccaria). Soil depth only influenced bacterial and fungal communities in barren and sparsely vegetated soils. These changes were partly due to more silt and higher soil moisture in the surface. In both soil ages, the surface was characterized by OTUs affiliated to Phormidium and Sphingobacteriales. In lower depths, however, bacterial and fungal communities differed between SSD. Lower depths of sparsely vegetated soils consisted of OTUs affiliated to Acidobacteria and Geoglossum, whereas depths of barren soils were characterized by OTUs related to Gemmatimonadetes. Overall, plant establishment drives the soil microbiota along the successional gradient but does not influence the vertical distribution of microbiota in recently deglaciated soils.     

Patchiness and disturbance: plant community responses to porcupine diggings in the central Negev

We tested the hypothesis that diversity and productivity of herbaceous plant communities in disturbed soil are related to the physical and biological heterogeneity of the landscape Our study was earned out on vegetation responses in porcupine diggings on a rocky slope in the central Negev desert in Israel We measured aboveground bio-mass and plant density per species in 150 porcupine diggings (15 cm deep and 15 to 20 cm wide) and in equally sized adjacent control samples in the undisturbed soil matrix We calculated mean annual biomass production, plant density and species richness for 10 sample areas along the slope In addition, we divided the plants into groups according to propagule size and dispersal mode We denoted two types of landscape heterogeneity, which we called physical and biological patchiness Physical patchiness was measured as the ratio of bare rock to soil surface Biological patchiness was the area of the soil covered by shrubs with associated soil mound and under-story relative to the total soil surface We also measured disturbance density, as the long term (17 yr) average density of newly made porcupine diggings We found that 1) the physical patchiness explained 30% of the variation of biological patchiness along the slope, while 2) the patterns of disturbance intensity and biological patchiness were similar (R-=0 386) 3) Biomass, density and species richness were significantly higher in diggings than m the soil matrix 4) Plant density in the matrix, but not m the diggings, was significantly correlated with physical patchiness, 5) species richness in diggings was significantly correlated with biological patchiness, but 6) biomass production in diggings and matrix was not affected by either physical or biological patchiness of the landscape 7) Disturbance density did not affect vegetation responses in diggings and matrix 8) A shift in the plant communities in the matrix towards plants with smaller seeds was associated with increasing physical patchiness, while m diggings there was an opposite shift 9) The proportion of wind dispersers was higher in diggings than outside, while the proportion of runoff dispersers was lower, 10) the densities of runoff dispersers in diggings and matrix were positively correlated with physical and biological patchiness 11) Physical and biological patchiness formed the two major gradients of species composition, explaining 30 and 25% respectively We conclude that the network of physical and biological patchiness and soil disturbance are important in the redistribution of resources and seeds, which control plant biomass, density, species richness and diversity The bare rock surface is the main source for runoff flow with associated soil, organic matter and nutrients The understory vegetation of shrubs provides seeds for creating and maintaining diversity The soil matrix absorbs runoff flow, and disturbances absorb runoff and trap seeds Thus, differences in landscape heterogeneity and their effects on resource and seed movement interact in controlling plant community productivity and diversity in the landscape     

Effects of environmental context on the susceptibility of Atriplex patula to attack by herbivorous beetles

The susceptibility of plants to attack by insect herbivores often depends on local environmental conditions. This study documents variation in herbivore damage by the chrysomelid beetle Erynephala maritima to the annual forb Atriplex patula in two microhabitats within New England salt marshes: bare patches and dense matrix vegetation. Environmental conditions within bare patches differ from those within matrix vegetation in a number of ways. Bare patches are characterized by the absence of perennial grasses and rushes (matrix vegetation) and greater levels of physical stress, and are rapidly colonized by the fugitive annual, Salicornia europaea, a second host plant of these beetles. Surveys of herbivore damage across three marshes revealed that A. patula in bare patches had a greater proportion of leaves damaged by beetles than those within matrix vegetation. Presence or absence of matrix vegetation and presence or absence of S. europaea were experimentally manipulated to determine the proximate cause of this pattern. The presence of S. europaea significantly increased the susceptibility of A. patula to herbivory in experimental plots. Both the extent of herbivore damage to plants and the proportion of plants damaged through time were greater in treatments with S. europaea than in controls, regardless of the presence or absence of matrix vegetation. Plants in S. europaea addition treatments were also less likely to survive to reproduction. Decreased survival appears to result from increased herbivory, suggesting that the negative effect of S. europaea on A. patula is mediated indirectly through shared insect herbivores. These results support the hypothesis that indirect interactions between alternative host plants, mediated by insect herbivores, can be important in natural communities. Received: 9 January 1999 / Accepted: 29 April 1999     

Influence of drought on plant performance through changes in belowground tritrophic interactions

Climate change is predicted to increase the risk of drought in many temperate agroecosystems. While the impact of drought on aboveground plant‐herbivore‐natural enemy interactions has been studied, little is known about its effects on belowground tritrophic interactions and root defense chemistry. We investigated the effects of low soil moisture on the interaction between maize, the western corn rootworm (WCR, Diabrotica virgifera), and soil‐borne natural enemies of WCR. In a manipulative field experiment, reduced soil moisture and WCR attack reduced plant performance and increased benzoxazinoid levels. The negative effects of WCR on cob dry weight and silk emergence were strongest at low moisture levels. Inoculation with entomopathogenic nematodes (EPNs, Heterorhabditis bacteriophora) was ineffective in controlling WCR, and the EPNs died rapidly in the warm and dry soil. However, ants of the species Solenopsis molesta invaded the experiment, were more abundant in WCR‐infested pots and predated WCR independently of soil moisture. Ant presence increased root and shoot biomass and was associated with attenuated moisture‐dependent effects of WCR on maize cob weight. Our study suggests that apart from directly reducing plant performance, drought can also increase the negative effects of root herbivores such as WCR. It furthermore identifies S. molesta as a natural enemy of WCR that can protect maize plants from the negative impact of herbivory under drought stress. Robust herbivore natural enemies may play an important role in buffering the impact of climate change on plant‐herbivore interactions.