Interacting coexistence mechanisms in annual plant communities: Frequency-dependent predation and the storage effect

We study frequency-dependent seed predation (FDP) in a model of competing annual plant species in a variable environment. The combination of a variable environment and competition leads to the storage-effect coexistence mechanism (SE), which is a leading hypothesis for coexistence of desert annual plants. However, seed predation in such systems demands attention to coexistence mechanisms associated with predation. FDP is one such mechanism, which promotes coexistence by shifting predation to more abundant plant species, facilitating the recovery of species perturbed to low density. When present together, FDP and SE interact, undermining each other’s effects. Predation weakens competition, and therefore weakens mechanisms associated with competition: here SE. However, the direct effect of FDP in promoting coexistence can compensate or more than compensate for this weakening of SE. On the other hand, the environmental variation necessary for SE weakens FDP. With high survival of dormant seeds, SE can be strong enough to compensate, or overcompensate, for the decline in FDP, provided predation is not too strong. Although FDP and SE may simultaneously contribute to coexistence, their combined effect is less than the sum of their separate effects, and is often less than the effect of the stronger mechanism when present alone.     

The role of specialist parasites in structuring host communities

Natural enemies can be a powerful force when structuring natural communities, and in facilitating or preventing species coexistence depending on the nature of the trophic interaction. In particular, “keystone” predators can promote species coexistence, provided they preferentially attack the competitively dominant species. However, it is not clear whether parasites can play a similar structuring role; parasites typically form chronic associations with their victims, reducing their fitness (i.e., fecundity) rather than survival, and allowing infected hosts to remain viable competitors within the community. Therefore the density-dependent suppression of the host is likely to be more subtle than that due to predation. Using a series of simple population-dynamic models we show that specialist parasites can facilitate species coexistence, although possibly less so than predators. These results contrast with those typically found with models of generalist parasites, which can reduce the likelihood of species coexistence through apparent competition. In addition, we show that the likelihood of parasite-facilitated species coexistence depends greatly on the specific type of parasite. In particular, macroparasites (e.g., parasitic helminths) may be less likely to facilitate species coexistence than microparasites (e.g., viruses or bacteria) due to their typically highly aggregated distribution amongst their hosts. Furthermore, species coexistence is more likely if the parasite is relatively benign to its host. Parasitism by apparently “harmless” specialist parasites may provide an important but overlooked factor in the maintenance of species diversity, facilitating species invasions into new communities and the emergence of novel infectious diseases.     

一年生植物群落内取样效应和互补效应的分离

利用人工建植的1年生植物群落对物种多样性．生产力关系及其潜在作用机理——取样效应和互补效应进行了研究。结果表明,不同多样性水平内群落生产力变化幅度较大,不同的物种组合对生产力水平也有较大的影响。植物种多样性与群落生产力之间呈二次函数关系。计算表明,所有混播群落均程度不同地存在多样性净效应,但不同多样性水平间及同一多样性水平内,净效应变化较大。物种多样性与生产力净效应间亦呈二次函数关系。多样性净效应与群落生产力水平呈显著正相关关系。多样性净效应的两个组成部分——取样效应和互补效应对多样性的变化有不同的反应：互补效应与物种多样性间呈单峰格局关系;而取样效应与多样性间呈显著的正相关关系。取样效应和互补效应均随着群落生产力的提高而显著增强。结果还表明,除物种多样性外,物种的组成（或物种的特性）对群落生产力、多样性净效应及其分解成分——取样效应和互补效应均有显著影响。据此,可以认为在短期多样性实验中,取样效应和互补效应均对多样性．生产力正相互关系产生作用,但取样效应的强度要大于互补效应。     

植物群落多样性与可入侵性关系研究进展

综述了植物群落的多样性和可入侵性关系的研究进展.对模型研究、观察性研究和实验性研究得出植物群落的多样性和可入侵性关系不同结论的原因进行了分析,并提出了该方面进一步研究的建议.模型研究的群落多样性形成机制单一、群落状态达到平衡、物种特征差异不明显等简化前提假设,削弱了其结论的可信性.观察性研究的尺度不同导致不同结论,在小尺度上的主要影响因素可能是群落内部的生物因素,但在较大尺度上很多外部因素共同影响多样性和可入侵性,会导致两者出现正相关.实验性研究通常是在小尺度下进行,排除了外部因素的干扰,但是在人工组建群落多样性梯度时,会出现取样效应、相同物种多样性的群落丰富度和密度不同等问题.未来研究应重点考虑的几个方面,即观察性研究与实验相结合,分析不同尺度上入侵的机理和过程;平衡实验设计,避免抽样效应等误来源;模型与实验设计相耦合.     

Variation in moisture duration as a driver of coexistence by the storage effect in desert annual plants

Temporal environmental variation is a leading hypothesis for the coexistence of desert annual plants. Environmental variation is hypothesized to cause species-specific patterns of variation in germination, which then generates the storage effect coexistence mechanism. However, it has never been shown how sufficient species differences in germination patterns for multispecies coexistence can arise from a shared fluctuating environment. Here we show that nonlinear germination responses to a single fluctuating physical environmental factor can lead to sufficient differences between species in germination pattern for the storage effect to yield coexistence of multiple species. We derive these nonlinear germination responses from experimental data on the effects of varying soil moisture duration. Although these nonlinearities lead to strong species asymmetries in germination patterns, the relative nonlinearity coexistence mechanism is minor compared with the storage effect. However, these asymmetries mean that the storage effect can be negative for some species, which then only persist in the face of interspecific competition through average fitness advantages. This work shows how a low dimensional physical environment can nevertheless stabilize multispecies coexistence when the species have different nonlinear responses to common conditions, as supported by our experimental data.     

Evidence of a component Allee effect driven by predispersal seed predation in a plant (Pedicularis rex,Orobanchaceae)

A small or sparse population may suffer a reduction in fitness owing to Allee effects. Here, we explored effects of plant density on pollination, reproduction and predation in the alpine herb Pedicularis rex over two years. We did not detect a significant difference in the pollination rate or fecundity (fruit set and the initial seed set) before predation between sparse and dense patches in either year, indicating no pollination-driven Allee effect. However, dense patches experienced significantly fewer attacks by predispersal seed predators in both years, resulting in a significantly decreased realized fecundity (final seed set), suggesting a component Allee effect driven by predispersal seed predation. Predation-driven Allee effects have been predicted by many models and demonstrated for a range of animals, but there is scant evidence for such effects in plants. Our study provides strong evidence of a component Allee effect driven by predation in a plant species.     

The roles of stochasticity and biotic interactions in the spatial patterning of plant species in alpine communities

Questions

Plant community composition can be influenced by multiple biotic, abiotic, and stochastic factors acting on the local species pool to determine their establishment success and abundance and subsequently the diversity of the community. We asked if the influences of biotic interactions on the composition of plant species in communities, as indicated by patterns of plant species spatial associations (independent, positive or negative), vary across a productivity gradient within a single ecosystem type. Do dominant species of communities show spatial patterning suggestive of competitive interactions with interspecific neighbors? Do species that span multiple community types exhibit the same heterospecific interactions with neighbours in each community?

Location

Three alpine communities in the southern Rocky Mountains.

Methods

We measured the occurrence of species in a 1‐cm spatial grid within 2 m × 2 m plots to determine the spatial patterns of species pairs in the three communities. A null model of independent species spatial arrangements was used to determine whether species pairs were positively, negatively or independently associated, and how these patterns differed among the communities across the gradient of resource supply and environmental stress.

Results

Positive associations, indicative of facilitation between species, were most common in the most resource‐poor and least productive community. However negative associations, suggestive of competitive interactions among species, were not more common in the two more resource‐rich, productive communities. The dominant species of these communities did exhibit higher negative than positive associations with neighbours relative to positive patterning. Independent interspecific patterning was equally common relative to positive and negative patterns in all communities. Species that previously were shown to either facilitate other species or compete with neighbours exhibited spatial patterning consistent with the earlier experimental work.

Conclusions

A large number of species exhibit a lack of net biotic interactions, and stochastic factors appear to be as important as competition and facilitation in shaping the structure of the three alpine plant communities we studied.

     

The contribution of nitrogen deposition to the eutrophication signal in understorey plant communities of European forests

We evaluated effects of atmospheric deposition of nitrogen on the composition of forest understorey vegetation both in space and time, using repeated data from the European wide monitoring program ICP‐Forests, which focuses on normally managed forest. Our aim was to assess whether both spatial and temporal effects of deposition can be detected by a multiple regression approach using data from managed forests over a relatively short time interval, in which changes in the tree layer are limited. To characterize the vegetation, we used indicators derived from cover percentages per species using multivariate statistics and indicators derived from the presence/absence, that is, species numbers and Ellenberg's indicator values. As explanatory variables, we used climate, altitude, tree species, stand age, and soil chemistry, besides deposition of nitrate, ammonia and sulfate. We analyzed the effects of abiotic conditions at a single point in time by canonical correspondence analysis and multiple regression. The relation between the change in vegetation and abiotic conditions was analyzed using redundancy analysis and multiple regression, for a subset of the plots that had both abiotic data and enough species to compute a mean Ellenberg N value per plot using a minimum of three species. Results showed that the spatial variation in the vegetation is mainly due to “traditional” factors such as soil type and climate, but a statistically significant part of the variation could be ascribed to atmospheric deposition of nitrate. The change in the vegetation over the past c. 10 years was also significantly correlated to nitrate deposition. Although the effect of deposition on the individual species could not be clearly defined, the effect on the vegetation as a whole was a shift toward nitrophytic species as witnessed by an increase in mean Ellenberg's indicator value.     

The effect of plant litter on ecosystem properties in a Mediterranean semi‐arid shrubland

Abstract. We studied the distribution of litter in a shrubland of the Negev with a semi‐arid Mediterranean climate of less than 200 mm of rainfall per year. Our focus was on the effects of litter on properties of landscape patches relevant to ecosystem processes (water runoff and soil erosion), annual plant community responses (seedling density, biomass production and species richness), and animal activity (soil disturbance by termites). Three 60‐m transects, extending across a pair of opposing north‐ and south‐facing slopes and their drainage channel, showed that litter accumulates not only under shrubs, but to a lesser extent also on the crusted inter‐shrub open areas. We used 35 experimental units (‘cells’, 0.5m × 1 m), each containing a crust and a shrub patch. Because runoff flows from crusted patches and is intercepted by shrub patches, the latter were in the lower third of the cells. Leaf litter was added in single and double amounts providing ca. 0.5 and 1.0 cm litter depth, to either, both, or none of the patches. Litter addition significantly decreased the amount of runoff, regardless of the location and amount of litter applied. Litter on the crust increased species number and seedling density of species with low abundance. Adding a double litter layer increased annual plant biomass production, while a single amount had no effect. Litter addition to the shrub patch affected neither biomass nor species richness. Litter addition to both patches at both quantities caused a large increase in termite activity. Termites caused disturbance by disrupting the crust, which may contribute to the reduction in runoff amounts. In the open, flat crust patches, annual plant communities are limited in their productivity and species richness, as there are few structures stopping the outflow of water, soil and seeds. Litter adds such structures, but affects the plant communities only when added to litter‐free crust. Litter accumulation and its patchy distribution have large impacts on landscape patch properties affecting resource distribution, plant productivity and diversity, and animal activity. Therefore, understanding litter distribution in relation to the patchy structure of the landscape of semi‐arid shrubland should be viewed as an important component of shrubland management.     

薇甘菊对香港郊野公园植物群落危害的分析

从薇甘菊(Mikania micrantha)对不同类型群落的危害,群落中各植物种类受薇甘菊危害的程度,以及薇甘菊与群落中主要乔木和灌木的种间联结性等方面,对香港郊野公园植物群落受薇甘菊危害状况进行了分析。研究样地所在的植被受薇甘菊的危害严重,草灌丛中薇甘菊的平均覆盖率高达80%,灌木群落中薇甘菊的平均覆盖率为60%左右,常绿阔叶林中的薇甘菊覆盖率平均约为30%。对香港郊野公园受薇甘菊危害的植物种类分析表明:受薇甘菊危害最大的主要是乔木和灌木,草本和藤本的受害比例和受害程度相对较小。受薇甘菊危害的乔木有30种,占群落中乔木的62.50%;受薇甘菊危害的灌木有29种,占群落中灌木的70.73%;受薇甘菊危害的草本植物有10种,占群落中草本植物的33.33%。通过无样方法采用2×2列联表的χ2检验对薇甘菊与群落中主要乔木和灌木的种间联结性分析表明:与薇甘菊呈正联结的乔木有12种,占主要乔木(28种)的42.86%,这些种大多是阳生性乔木;与薇甘菊呈负联结的乔木有16种,占主要乔木的57.16%。与薇甘菊呈正联结的灌木有11种,占主要灌木(19种)的57.89%,多是阳生性灌木;与薇甘菊呈负联结的灌木有8种,占主要灌木的42.11%,这些种大多数也是阳生性灌木,仅九节是耐阴性灌木。     

Selection on plant traits in hedgerow ground vegetation: The effect of time since conversion from conventional to organic farming

This paper aims at characterizing the response of hedgerow ground flora to conversion of the surrounding fields from conventional farming to an organic farming practice. The effect of time since conversion from conventional to organic farming on the neighbouring hedgerow vegetation was examined by measuring selection on plant species and plant traits in the ground flora of 57 hedgerows adjacent to agricultural fields subject to organic or conventional management. Species richness increased significantly with time since conversion to organic farming until a maximum was reached. There were significant changes in community weighted average trait value in seven out of fifteen plant traits, and in functional diversity in two out of three groups of traits. In contrast to hypotheses, transition to organic farming generally selected for plant species that are less adapted to competing for light, whereas there were no effects on life form and lifespan. When all plant traits were analyzed together in a regression tree analysis, 31% of the variance was explained by five plant traits in the pruned tree, indicating that factors other than farming practice have a large influence on hedgerow ground flora.     

The whole and its parts: why and how to disentangle plant communities and synusiae in vegetation classification

Most plant communities consist of different structural and ecological subsets, ranging from cryptogams to different tree layers. The completeness and approach with which these subsets are sampled have implications for vegetation classification. Non‐vascular plants are often omitted or sometimes treated separately, referring to their assemblages as “synusiae” (e.g. epiphytes on bark, saxicolous species on rocks). The distinction of complete plant communities (phytocoenoses or holocoenoses) from their parts (synusiae or merocoenoses) is crucial to avoid logical problems and inconsistencies in the resulting classification systems. We here describe theoretical differences between the phytocoenosis as a whole and its parts, and outline consequences of this distinction for practice and terminology in vegetation classification. To implement a clearer separation, we call for modifications of the International Code of Phytosociological Nomenclature and the EuroVegChecklist. We believe that these steps will make vegetation classification systems better applicable and raise recognition of the importance of non‐vascular plants in the vegetation as well as their interplay with vascular plants.