Influence of temporal heterogeneity in nitrogen supply on competitive interactions in a desert shrub community

Soil nutrients in arid systems are supplied to plants in brief pulses following precipitation inputs. While these resource dynamics have been well documented, little is known about how this temporal heterogeneity influences competitive interactions. We examined the impacts of the temporal pattern of N supply on competitive intensity and ability in an N-limited desert shrub community. At our field site, the three codominant shrubs, Atriplex confertifolia, A. parryi, and Sarcobatus vermiculatus, differ in seasonal growth patterns, with A. confertifolia and S. vermiculatus achieving higher growth rates earlier in the growing season than A. parryi. We predicted that these timing differences in maximum growth rate may interact with temporal variation in N supply to alter competitive abilities over time. Seedlings of the two Atriplex species were planted either individually in field plots or as target plants surrounded by neighbor seedlings. After one year of establishment, the same amount of ¹⁵N was applied to plots either as early spring pulses, mid spring pulses or continuously through the second growing season. Competitive effects were observed under continuous and pulsed N supply. Averaged across all target–neighbor treatments, competitive intensity was ∼1.8-fold greater when N was pulsed compared to when N was supplied continuously, but overall, the outcome of competitive interactions was not influenced by N pulse timing. While the timing of resource supply did not differentially influence the competitive abilities of coexisting species in this system, the temporal pattern of resource supply did alter the intensity of competitive interactions among species. While additional studies in other systems are needed to evaluate the generality of these results, this study suggests that competitive intensity may not necessarily be a direct function of productivity or resource availability as traditionally assumed. Instead, the intensity of competitive interactions in resource-poor systems may depend upon the temporal pattern of resource supply.     

Root system size determines plant performance following short-term soil nutrient pulses

Heterogeneity in soil resources is common in natural communities, in both space and time. In contrast to substantial research into plant responses to spatial nutrient heterogeneity, little is known about the factors that influence plants responses to temporally heterogeneous nutrient supplies. We examine (1) the effects of nutrient pulse length on plant biomass allocation patterns, (2) whether a plant??s root system prior to a nutrient pulse influences the benefit a plant can gain from that pulse, and (3) how competition and nutrient pulses interact to influence plant biomass. To address these issues, we used a pot experiment with varying initial conditions (nutrient levels and competition) designed to produce a range of root system sizes in the grass Poa pratensis. We then subjected the plants to short-term (3 and 10?days long) nutrient pulses where the same total nutrients were delivered at varying rates per unit time (intensity). Plant biomass and tissue nitrogen concentration were lowest when pulses were short relative to either long pulses or continuous supply. Plants with larger root systems at the start of the nutrient pulse gained more benefit than smaller plants because final plant size relative to initial root system size decreased as pulse intensity (nutrients delivered per unit time) increased. This study has two important implications. First, since larger plants benefit disproportionately more from a nutrient pulse than smaller plants, pre-pulse conditions such as the competitive environment and resource supply are likely key determinants of a plant??s ability to capture nutrients from a pulse. Second, the importance of pre-pulse size provides a belowground mechanism to enhance the size variability among plants in a community.     

Competitive impacts and responses of an invasive weed: dependencies on nitrogen and phosphorus availability

Changes in competitive interactions under conditions of enhanced resource availability could explain the invasion success of some problematic plant species. For one invader of North American grasslands, Centaurea diffusa (diffuse knapweed), we test three hypotheses: (1) under ambient (high resource) conditions, C. diffusa is better able to tolerate competition from the resident community (competitive response), (2) under ambient conditions, C. diffusa strong impacts the competitive environment (competitive effect), and (3) reductions in nitrogen and/or phosphorus availability diminish these advantages. In support of our first hypothesis, C. diffusa was the most tolerant to neighbor competition of the four focal species under current resource conditions. In opposition to our second hypothesis, however, neighborhoods that contained C. diffusa and those where C. diffusa had been selectively removed did not differ in their impact on the performance of target transplant individuals or on resource conditions. Reduction in resource availability influenced competitive tolerance but not competitive impact, in partial support of our last hypothesis. Reduction in soil nitrogen (via sucrose carbon addition) enhanced the degree of neighbor competition experienced by all species but did not change their relative rankings; C. diffusa remained the best competitor under low nitrogen conditions. Reduction of soil phosphorus (via gypsum addition) weakened the ability of C. diffusa to tolerate neighbor competition proportionately more than the other focal species. Consequently, under low phosphorus conditions, C. diffusa lost its competitive advantage and tolerated neighbor competition similarly to the other focal species. We conclude that C. diffusa invasion may be double-edged: C. diffusa is less limited by nitrogen than the other focal species and is better able to utilize phosphorus to its competitive advantage.     

Effects of neighbouring vegetation on eucalypt seedlings at two sites subject to different levels of abiotic stress

Abstract In spite of numerous studies on the effect of nutrient levels and/or standing crop on the intensity of resource competition the debate has not been resolved. Field studies that have used natural productivity gradients have generally supported the argument that competitive intensity and resource availability are positively correlated, whereas studies that have used artificial resource gradients have generally refuted the same argument. Here we report the results from study in which both approaches were used within the same system. We studied two species of eucalypt that occupy contrasting parts of the same landscape: Eucalyptus camaldulensis, found mostly along creek lines and in valleys with deep alluvial soils, and Eucalyptus microcarpa, found on hillsides and ridges with shallow soils. We studied the response of seedlings of the two species to the combined effects of competition and manipulated nutrient levels in a glasshouse experiment, and also investigated their responses to removal of neighbouring plants in the field. Eucalyptus microcarpa was less responsive to increased resource availability, which is consistent with one of the principal assumptions of Grime’s C‐S‐R model. In the glasshouse experiment both species of eucalypt responded in a qualitatively similar fashion to the combined effects of resource availability and competition: release from competition resulted in increased growth, but only in pots that received additional resources. In the field we found that neighbouring vegetation could severely limit the establishment of E. camaldulensis but the removal of neighbouring vegetation did not affect the performance of E. microcarpa seedlings. Eucalyptus camaldulensis seedlings suffered high levels of damage from herbivores. Our results thus generally support the predications of the C‐S‐R model, however, they indicate that the effects of competition and herbivory may be heavily confounded.     

养分资源脉冲供给对几种微藻种间竞争的影响

以淡水生态系统常见的5种微藻为研究对象,通过稳态条件下单一培养的方式获取各微藻在氮素或磷素缺乏条件下对应的生长特征参数和R*值,同时将5种微藻在养分资源脉冲供给的方式下进行混合培养,进而检测养分资源脉冲供给对几种微藻种间竞争的影响作用,并与基于稳态条件下所预测的微藻种间竞争结果进行比较。研究结果显示,在稳态条件下具有最小R*值的纤细角星鼓藻在与其它微藻的种间竞争过程中始终保持优势地位,而其余4种微藻在混合培养状态下的相对比重亦与其稳态条件下的R*值紧密相关。然而,资源脉冲供给条件下的竞争优势种纤细角星鼓藻能与其它种群数量较小的微藻共存。此外,在氮素和磷素的不同供给比例情况下,对应微藻群落的结构也会发生相应的变化。实验养分资源脉冲作用下多种微藻的共存现象与自然水体中的观测现象相一致,显示了资源脉冲可能是维持生物多样性水平的一个重要机制。     

Competition for pulsed resources: an experimental study of establishment and coexistence for an arid-land grass

In arid environments, episodically-pulsed resources are important components of annual water and nutrient supply for plants. This study set out to test whether seedlings have an increased capacity for using pulsed resources, which might then improve establishment when in competition with older individuals. A second aim was to determine whether there is a trade-off in competitive strategies when resources are supplied continuously at low concentrations, or as pulses with pronounced inter-pulse periods. A glasshouse experiment used a target-neighbour design of size-asymmetric competition, with juveniles of Panicum antidotale (blue panicgrass) introduced into contrasting densities of adult plants. Stable isotopes of nitrogen were used for measuring plant resource uptake from pulses, and tolerance to inter-pulse conditions was assessed as the mean residence time (MRT) of nitrogen. A higher root/shoot ratio and finer root system enhanced the capacity of juveniles to use resources when pulsed, rather than when continuously supplied. Higher resource uptake during pulses improved the establishment of juvenile Panicum in mixed cultures with older individuals. However, a trade-off was observed in plant strategies, with juveniles showing a lower MRT for nitrogen, which suggested reduced tolerance to resource deficit during inter-pulse periods. Under field conditions, higher utilization of pulsed resources would lead to the improved seedling establishment of Panicum adjacent to “nurse” plants, whereas mature plants with well-developed roots, exploiting a greater soil volume, maintain more constant resource uptake and retention during inter-pulse periods.     

Precipitation pulse use by an invasive woody legume: the role of soil texture and pulse size

Plant metabolic activity in arid and semi-arid environments is largely tied to episodic precipitation events or “pulses”. The ability of plants to take up and utilize rain pulses during the growing season in these water-limited ecosystems is determined in part by pulse timing, intensity and amount, and by hydrological properties of the soil that translate precipitation into plant-available soil moisture. We assessed the sensitivity of an invasive woody plant, velvet mesquite (Prosopis velutina Woot.), to large (35 mm) and small (10 mm) isotopically labeled irrigation pulses on two contrasting soil textures (sandy-loam vs. loamy-clay) in semi-desert grassland in southeastern Arizona, USA. Predawn leaf water potential (Ψ_pd), the isotopic abundance of deuterium in stem water (δD), the abundance of ¹³C in soluble leaf sugar (δ¹³C), and percent volumetric soil water content (θ_v) were measured prior to irrigation and repeatedly for 2 weeks following irrigation. Plant water potential and the percent of pulse water present in the stem xylem indicated that although mesquite trees on both coarse- and fine-textured soils quickly responded to the large irrigation pulse, the magnitude and duration of this response substantially differed between soil textures. After reaching a maximum 4 days after the irrigation, the fraction of pulse water in stem xylem decreased more rapidly on the loamy-clay soil than the sandy-loam soil. Similarly, on both soil textures mesquite significantly responded to the 10-mm pulse. However, the magnitude of this response was substantially greater for mesquite on the sandy-loam soil compared to loamy-clay soil. The relationship between Ψ_pd and δ¹³C of leaf-soluble carbohydrates over the pulse period did not differ between plants at the two sites, indicating that differences in photosynthetic response of mesquite trees to the moisture pulses was a function of soil water availability within the rooting zone rather than differences in plant biochemical or physiological constraints. Patterns of resource acquisition by mesquite during the dynamic wetting–drying cycle following rainfall pulses is controlled by a complex interaction between pulse size and soil hydraulic properties. A better understanding of how this interaction affects plant water availability and photosynthetic response is needed to predict how grassland structure and function will respond to climate change.     

丛枝菌根对紫茎泽兰和黄花蒿地上地下养分分配的竞争调控策略

紫茎泽兰（Eupatorium adenophorum）入侵喀斯特生态系统导致群落多样性和稳定性降低是该区域面临的重要生态问题,丛枝菌根（Arbuscular mycorrhizae,AM）通过根系外延菌丝互联不同物种个体影响植物养分竞争,但如何调控入侵种与乡土种地上地下资源竞争分配尚不清楚。以入侵种紫茎泽兰和乡土种黄花蒿（Artemisia annua）为研究对象,使用由1个竞争室和2个种植室所组成的微生态系装置。针对两物种对竞争室养分资源利用,采用20 μm和0.45 μm尼龙网设置共同竞争（CC）、单一利用（SU）和对照（CK）处理,并对上述处理进行AM真菌接种（M⁺）与不接种（M^-）,分析不同处理下紫茎泽兰与黄花蒿地下地上生物量及氮磷养分分配。结果表明：就地上部分而言,比较M⁺与M^-,三种竞争方式的紫茎泽兰磷吸收量均显著表现为M⁺ > M^-,但黄花蒿在M⁺与M^-间无显著差异;比较3种竞争方式,M⁺下紫茎泽兰氮吸收量和黄花蒿生物量及氮磷吸收量表现为SU>CK,黄花蒿地上生物量、氮磷吸收量则表现为CCSU;M⁺下CC处理紫茎泽兰地上氮吸收量显著高于黄花蒿;M^-下黄花蒿氮磷吸收量显著表现为CC和SU>CK,但紫茎泽兰在CC、SU和CK间无显著差异。就地下部分而言,三种竞争方式的紫茎泽兰地下生物量、氮磷吸收量显著表现为M⁺ > M^-,但黄花蒿在M⁺与M^-间无显著差异;比较三种竞争方式,M⁺下紫茎泽兰氮吸收量在CC与SU间无显著差异,但SU>CK;比较植物间,M⁺条件下,CC和SU处理的紫茎泽兰氮磷吸收量均显著高于黄花蒿,而M^-条件下紫茎泽兰与黄花蒿生物量和氮磷吸收量在三种竞争方式间均无显著差异。研究表明,AM真菌通过菌根网络调控入侵种和乡土种的竞争能力,影响公用土壤养分资源在植株地上地下的资源分配并提高入侵植物从菌根共生体中获得收益促进其入侵。     

Growth rates of rhizosphere microorganisms depend on competitive abilities of plants and N supply

Abstract

Plant‐microbial interactions under N‐limiting conditions are governed by competitive abilities of plants for N. Our study aimed to examine how two plant species of strawberry, Fragaria vesca L. (native species) and Duchesnea indica (Andrews) Focke (an invasive plant in central Europe), growing in intra‐specific and inter‐specific competition alter the functions of rhizosphere microorganisms in dependence on N availability. By intra‐specific competition at low N level, a 2.4‐fold slower microbial‐specific growth rate was observed under D. indica characterized by smaller root biomass and lower N content in roots compared with F. vesca. By inter‐specific competition of both plants at low N level, microbial growth rates were similar to those for D. indica indicating that plants with stronger competitive abilities for N controls microbial community in the rhizosphere. Since a high N level smoothed the differences between plant species in root and microbial biomass as well as in microbial growth rates under both intra‐specific and inter‐specific competition, we conclude that competitive abilities of plant species were crucial for microbial growth in the rhizosphere only under N imitation.     

To compete or not to compete: an experimental study of interactions between plant species with contrasting root behaviour

Game-theoretic models predict that plants with root systems that avoid belowground competition will be displaced by plants that overproduce roots in substrate shared with competitors. Despite this, both types of root response to neighbours have been documented. We used two co-occurring clonal species (Glechoma hederacea and Fragaria vesca) with contrasting root responses to neighbours (avoidance of competition and contesting of resources, respectively) to examine whether functional variation in other traits affected the success of each rooting strategy, leading to a different outcome from that predicted on the basis of root behaviour alone. Vegetative propagation rates, morphology and biomass allocation patterns were examined when each species was challenged with competition from physically separate ramets with either the same rooting strategy (intraclonal competition) or the contrasting rooting strategy (interspecific competition). Contrary to the predictions of game-theoretic models, the species that exhibits avoidance of root competition (Glechoma) was not competitively inferior to the species that does not (Fragaria). Glechoma achieved greater total mass in the interspecific treatment than in the intraclonal treatment. However, Fragaria did not experience more intense competition from Glechoma than it did in the intraclonal treatment. Strong interference between the two species appeared to be avoided because Glechoma invested preferentially in rapid exploitation of unoccupied space, whereas Fragaria invested in increasing the competitive ability and local persistence of established ramets. Our results suggest that interspecific trade-offs between traits related to competitive ability and resource exploitation can allow coexistence of species with contrasting rooting behaviours. Full assessment of the adaptive value of different root responses to neighbours therefore requires concurrent consideration of the combined effects of a wide array of functional traits.     

Nitrogen addition increases intraspecific competition in the invasive wetland plant Alternanthera philoxeroides,but not in its native congener Alternanthera sessilis

Nitrogen is often released in pulses with different frequencies, and N supply pulses may affect growth, reproduction, and biomass allocation of plants. However, few studies have examined how N supply pulses affect intraspecific competition of clonal plants and whether such an effect depends on the N supply amount. We grew one (no competition) or 12 ramets (with intraspecific competition) of both an invasive clonal plant Alternanthera philoxeroides and its native congener Alternanthera sessilis in five different N treatments: control (no N addition), low/high amount with low/high frequencies (pulses). Nitrogen addition significantly increased the growth of both species, while intraspecific competition decreased it. Nitrogen addition significantly increased intraspecific competitive intensity of A. philoxeroides as measured by the log response ratio of growth traits, but did not affect that of A. sessilis. Despite the N supply amount, N pulses had little effect on the growth and thus intraspecific competition of the two species. Therefore, increasing N deposition may change population structure and dynamics and the invasion succession of A. philoxeroides, but changes in N pulses may not.     

矮嵩草草甸主要植物不同器官对氮素的吸收及分配特征研究

以高寒矮嵩草(Kobresia humilis)草甸7个主要植物种为研究对象,利用15 N同位素标记技术,通过分析不同器官对氮素的吸收及分配特征,揭示主要植物种在群落中的生态适应性、竞争力和地位。结果显示:(1)矮嵩草的叶和茎、垂穗披碱草(Elymus nutans)的叶,以及双柱头藨草(Scirpus distigmaticus)和鹅绒委陵菜(Potentilla anserina)的叶、茎、根均偏好累积硝态氮,早熟禾(Poa annua)的穗和叶以及甘肃马先蒿(Pedicularis kansuensis)和短穗兔耳草(Lagotis brachystachya)的根均偏好积累铵态氮。(2)矮嵩草对吸收的甘氨酸和硝态氮主要分配于叶中,铵态氮分配于茎中;双柱头藨草对吸收的甘氨酸和硝态氮主要分配于茎中,铵态氮分配于叶中;垂穗披碱草和早熟禾对吸收的硝态氮和铵态氮主要分配于叶中;垂穗披碱草对吸收的甘氨酸主要分配于根中,而早熟禾将较多的甘氨酸分配到穗中;甘肃马先蒿对吸收的硝态氮主要分配于叶中,铵态氮分配于根中;鹅绒委陵菜对吸收的甘氨酸、硝态氮和铵态氮主要分配于叶中;短穗兔耳草对吸收的甘氨酸主要分配于叶中,硝态氮和铵态氮主要分配于根中。(3)在牧草生长盛期,矮嵩草草甸土壤的有机氮和无机氮主要贡献于甘肃马先蒿的花、早熟禾的穗、垂穗披碱草的根和鹅绒委陵菜的茎叶。研究表明,高寒矮嵩草草甸主要植物不同器官对氮素的吸收及分配呈现多元化特征,因不同植物种的生物学特性和生态适应习性而异。     

Does fluctuating resource availability increase invasibility? Evidence from field experiments in New Zealand short tussock grassland

The theory of fluctuating resource availability proposes that the susceptibility of a plant community to invasion by new species (i.e., invasibility) depends upon conditions of intermittent resource enrichment coinciding with the presence of invading propagules. We compared the response of a rapidly invading forb (Hieracium pilosella L.) between different experimental treatments in a short tussock grassland in New Zealand, over 6–12 years, to determine whether the theory explains differences in invasibility. The theory predicts that environments subject to periodic resource enrichment will be more invasible than those with more stable resource-supply rates. In our study, H. pilosella did not increase more rapidly in treatments subject to periodic resource pulses (fertiliser and water) than in those with more stable resource supplies. Also contrary to the predictions of the theory, the rate of invasion of H. pilosella did not increase following an increase in the rate of supply of water or nutrient resources, or following treatments that temporarily reduced resource uptake in the community, including grazing. H. pilosella did not increase immediately following abrupt increases in water and nutrient supply and removal of the dominant grass species with herbicide, as predicted by the theory, although temporary increases in resident exotic guilds indicated that the intensity of competition for resources was reduced. Neither H. pilosella nor resident exotic guilds showed increased cover growth rates following resumed grazing. The rate of invasion by H. pilosella was not correlated with species richness, a result consistent with one of the predictions of the theory. Therefore, short-lived events that temporarily reduced or suspended competition did not appear to determine the invasion success of this particular species in this region. In New Zealands perennial short tussock grasslands, the characteristics of the resident plant community may be more critical than resource fluctuations in determining invasion success of H. pilosella. Invasion of H. pilosella may be most successfully controlled here by promoting a successional physiognomic shift to a taller, shrub-and-tussock-dominated canopy that competitively excludes low-growing forbs.     

Additive effects of aboveground and belowground herbivores on the dominance of spotted knapweed (Centaurea stoebe)

Spotted knapweed (Centaurea stoebe) is found in over 3 million ha of rangeland and forests across North America, and evidence supporting the use of biological control as a regional method to reduce infestations and their associated impacts remains inconclusive. Several species of insects have been reported to reduce plant densities in some areas; however, rigorous studies that test combinations of these species and the influence of resource availability are lacking. We examined the singular and combined effects of herbivory by a root weevil (Cyphocleonus achates) and a flower head weevil (Larinus minutus) on the growth and flower production of C. stoebe. We also manipulated soil resource fertility as an additional factor that could explain the outcomes of contradictory biological control herbivore effects on C. stoebe. In a greenhouse study, herbivory by C. achates decreased flower production for plants across all resource environments. In a caged common garden study, the negative effects of herbivory also did not interact with soil nutrient status. However, the presence of plant competition further decreased knapweed growth, and the negative effects of concurrent herbivory by C. achates and L. minutus on plant biomass and flower production were additive. Derived within the context of variable levels of soil nutrient availability and competing vegetation, these results support the cumulative stress hypothesis and the contention that combined above- and belowground herbivory can reduce spotted knapweed densities and reduce the ecological and economic impacts of this species in rangelands of western North America.     

Effect of competition intensity on recruitment of palatable and unpalatable grasses

In this study, we made an attempt to reveal how competition intensity from established plants impacts on palatable and unpalatable grass seedlings recruitment, in a natural mesic grassland of central Argentina. Our objective was to assess the seedling recruitment of a palatable species (Chascolytrum subaristatum) and an unpalatable species (Nassella trichotoma) in microsites differing in competition intensity from established plants. Identity (C. subaristatum and N. trichotoma) and defoliation severity were used as surrogate for competition intensity. In March 2017, we permanently marked established individuals of N. trichotoma and C. subaristatum and placed two circular plots adjacent to each individual. In one plot we added seeds of N. trichotoma and in the other seeds of C. subaristatum. After seeding, established plants were randomly assigned to one of three level of defoliation: without defoliation, low defoliation severity and high defoliation severity. From April to November 2017 (i.e. over a complete annual growing cycle), we measured seedling density, recruitment and growth. Our results supported the hypothesis that seedlings of palatable grasses are more competitive than seedlings of unpalatable grasses. Seedling of the palatable grass C. subaristatum recruited successfully regardless the intensity of competition from established plants, whereas seedlings of the unpalatable grass N. trichotoma recruited better under low competitive pressure from established plants. Our results suggest that the availability of microsites with low competitive pressure from the established vegetation, created by selective grazing of palatable grasses, promotes the recruitment of unpalatable grass seedlings. This mechanism may contribute to the species replacement process commonly observed in heavy grazed grasslands.     

施氮对空心莲子草(Alternanthera philoxeroides)和莲子草(Alternanthera sessilis)种间关系的影响

自然界的氮素释放总是呈现出空间和时间上的异质性,但关于异质性氮释放对于入侵植物和本地植物种间关系影响的研究相对较少。将入侵植物空心莲子草(Alternanthera philoxeroides)和同属本地植物莲子草(Alternanthera sessilis)分别进行单种种植(12株,无种间竞争)和混种种植(每种6株,有种间竞争),模拟大气氮湿沉降设置由两种不同施氮总量(15g N m~(-2)a~(-1)和30g N m~(-2)a~(-1))和两种不同施氮频率(每5天1次和每15天1次)交叉组成的4种施氮处理,并以不施氮为对照。施氮总量的增加显著促进了两种植物的生长,但对两种植物的种间竞争关系没有显著影响。施氮频率对两种植物的生长以及种间竞争关系都没有显著影响。两种植物在面对竞争时表现出不同的生物量分配策略,空心莲子草将更多的生物量分配到茎,而莲子草将更多的生物量分配到根。在全球变化的背景下,大气氮湿沉降可能会改变两种植物的种群结构和动态,但可能对这两种植物的种间关系影响较小。     

Links between the detritivore and the herbivore system: effects of earthworms and Collembola on plant growth and aphid development

Effects of Collembola (Heteromurus nitidus and Onychiurus scotarius) and earthworms (Aporrectodea caliginosa and Octolasion tyrtaeum) on the growth of two plant species from different functional groups (Poa annua and Trifolium repens), and on the development of aphids (Myzus persicae) were investigated in a laboratory experiment lasting 20 weeks. Using soil from a fallow site which had been set aside for about 15 years, we expected that nitrogen would be of limited supply to plants and hypothesized that the soil animals studied, particularly earthworms, would increase nutrient availability to plants and thereby also modify aphid reproduction and development. Plant growth was modified strongly by the presence of soil animals. Earthworms caused a more than twofold increase in shoot and root mass of P. annua but increased that of T. repens by only 18% and 6%, respectively. However, earthworms neither affected plant shoot/root ratio nor the nitrogen concentration in plant tissue. In contrast, the presence of Collembola caused a reduction in plant biomass particularly that of P. annua roots, but plant tissue nitrogen concentration was increased, although only slightly. Aphid reproduction on T. repens was lowered in the presence of Collembola on average by 45% but on P. annua increased by a factor of about 3. It is concluded that Collembola decrease aphid reproduction on more palatable host plants like T. repens but increase that on less palatable ones like P. annua. Earthworm presence also affected aphid reproduction but the effect was less consistent than that of Collembola. In the presence of earthworms, aphid reproduction was in one experimental period increased by some 70%. Earthworms also modified the numbers of Collembola and their vertical distribution in experimental chambers. Exploitation of deeper soil layers by H. nitidus was increased but, generally, O. scotarius numbers were reduced whereas those of H. nitidus increased in earthworm treatments. The presence of Collembola also influenced earthworm body mass during the experiment. In general it declined, but in the presence of Collembola loss of body mass of A. caliginosa was more pronounced. We conclude that inhibiting effects between Collembola and earthworms resulted from the use of a common resource, litter material rich in nitrogen. This is supported by the higher C/N ratio of the litter material in the presence of earthworms and Collembola by the end of the experiment. Effects of soil invertebrates like Collembola and earthworms on plant performance and aphid development are assumed to be modified by complex direct and indirect interactions among soil animal groups. Received: 6 July 1998 / Accepted: 1 March 1999     

Dynamics of biomass partitioning in two competing meadow plant species

The optimal allocation theory predicts that growth is allocated between the shoot and the roots so that the uptake of the most limiting resource is increased. Allocation is dynamic due to resource depletion, interaction with competitors, and the allometry of growth. We assessed the effects of intra- and inter-specific competition on growth and resource allocation of the meadow species Ranunculus acris and Agrostis capillaris, grown in environments with high (+) or low (−) availability of light (L) and nutrients (N). We took samples twice a week over the 7 weeks experiment, to follow the changes in root-to-shoot ratios in plants of different sizes, and carried out a larger scale harvest at the end of the experiment. Of all the tested factors, availability of nutrients had the largest effect on the growth rate and shoot-to-root allocation in both species, although both competition and light had significant effects as well. The highest root-to-shoot ratios were measured from the L+N− treatment, and the lowest from the L−N+ treatment, as predicted by the optimal allocation theory. Competition changed resource allocation, but not always toward acquiring the resource that is most limiting to growth. We thus conclude that the greatest variation in shoot-to-root allocation was due to the resource availability and the effects of competition were small, probably due to low density of plants in the experiment.     

Effects of Nitrogen and Salinity on Growth and Competition Between a Native Grass and an Invasive Congener

Numerous studies show that an increase in the availability of limiting resources can increase invasion by non-native plants into natural communities. One possible explanation is that the ability of natives to compete with non-natives tends to decrease when resource availability is increased. We tested this hypothesis in a competition experiment using two closely matched plant species and two environmental factors related to limiting resources in a coastal grassland system on Bodega Head in northern California. We grew the native grass Bromus carinatus and the non-native grass B. diandrus together and apart at different levels of soil nitrogen crossed with different levels of soil salinity. Both species are abundant in the grassland and previous work suggested that the abundance of B. carinatus is lower and the abundance of B. diandrus is higher on soil that has been enriched with nitrogen. Salinity has been shown to be negatively associated with invasion by B. diandrus into another California grassland, and to vary significantly over short distances in the grassland at Bodega Head, where it could affect water availability, which strongly limits plant growth during the dry season. Contrary to our prediction that low resource availabilities would increase the relative competitive ability of the native, the ability of B. carinatus to compete with B. diandrus was not greater when nitrogen availability was lower or when soil salinity was higher. Instead, high salinity increased the relative competitive ability of the non-native, and low nitrogen had little effect on competition. This suggests that preventing resource enrichment will not suffice to control invasion by non-native plant species in this grassland.     

The shift from plant–plant facilitation to competition under severe water deficit is spatially explicit

The stress‐gradient hypothesis predicts a higher frequency of facilitative interactions as resource limitation increases. Under severe resource limitation, it has been suggested that facilitation may revert to competition, and identifying the presence as well as determining the magnitude of this shift is important for predicting the effect of climate change on biodiversity and plant community dynamics. In this study, we perform a meta‐analysis to compare temporal differences of species diversity and productivity under a nurse plant (Retama sphaerocarpa) with varying annual rainfall quantity to test the effect of water limitation on facilitation. Furthermore, we assess spatial differences in the herbaceous community under nurse plants in situ during a year with below‐average rainfall. We found evidence that severe rainfall deficit reduced species diversity and plant productivity under nurse plants relative to open areas. Our results indicate that the switch from facilitation to competition in response to rainfall quantity is nonlinear. The magnitude of this switch depended on the aspect around the nurse plant. Hotter south aspects under nurse plants resulted in negative effects on beneficiary species, while the north aspect still showed facilitation. Combined, these results emphasize the importance of spatial heterogeneity under nurse plants for mediating species loss under reduced precipitation, as predicted by future climate change scenarios. However, the decreased water availability expected under climate change will likely reduce overall facilitation and limit the role of nurse plants as refugia, amplifying biodiversity loss.