Rooting theories of plant community ecology in microbial interactions

Predominant frameworks for understanding plant ecology have an aboveground bias that neglects soil micro-organisms. This is inconsistent with recent work illustrating the importance of soil microbes in terrestrial ecology. Microbial effects have been incorporated into plant community dynamics using ideas of niche modification and plant–soil community feedbacks. Here, we expand and integrate qualitative conceptual models of plant niche and feedback to explore implications of microbial interactions for understanding plant community ecology. At the same time we review the empirical evidence for these processes. We also consider common mycorrhizal networks, and propose that these are best interpreted within the feedback framework. Finally, we apply our integrated model of niche and feedback to understanding plant coexistence, monodominance and invasion ecology.

The impact of Solanum elaeagnifolium, an invasive plant in the Mediterranean, on the flower visitation and seed set of the native co-flowering species Glaucium flavum

We examined the effect of the invasive Solanum elaeagnifolium (Solanaceae) on flower visitation patterns and seed set of the co-flowering native Glaucium flavum (Papaveraceae). We observed flowering G. flavum plants in invaded and uninvaded sites and found that G. flavum flowers in uninvaded sites received significantly more total visits. In addition, we hand-pollinated flowers on plants of G. flavum with (i) pure conspecific pollen, (ii) pure S. elaeagnifolium pollen and (iii) three different mixtures of the two types of pollen (containing 25, 50 and 75% invasive pollen). As a control, flowers were left unmanipulated or were permanently bagged. Seed set did not differ significantly between flowers receiving pollen mixtures and pure conspecific pollen. However, in the open pollination treatment, seed set was significantly lower than in the 100% conspecific pollen treatment, which suggests pollen limitation. Bagged flowers had very low seed set. G. flavum was generally resilient against the deposition of S. elaeagnifolium pollen.

A GIS-based empirical model for vegetation prediction in Lefka Ori, Crete

The aim of the study was to establish and verify a predictive vegetation model for plant community distribution in the alti-Mediterranean zone of the Lefka Ori massif, western Crete. Based on previous work three variables were identified as significant determinants of plant community distribution, namely altitude, slope angle and geomorphic landform. The response of four community types against these variables was tested using classification trees analysis in order to model community type occurrence. V-fold cross-validation plots were used to determine the length of the best fitting tree. The final 9node tree selected, classified correctly 92.5% of the samples. The results were used to provide decision rules for the construction of a spatial model for each community type. The model was implemented within a Geographical Information System (GIS) to predict the distribution of each community type in the study site. The evaluation of the model in the field using an error matrix gave an overall accuracy of 71%. The user's accuracy was higher for the Crepis-Cirsium (100%) and Telephium-Herniaria community type (66.7%) and relatively lower for the Peucedanum-Alyssum and Dianthus-Lomelosia community types (63.2% and 62.5%, respectively). Misclassification and field validation points to the need for improved geomorphological mapping and suggests the presence of transitional communities between existing community types.

A GIS-based empirical model for vegetation prediction in Lefka Ori, Crete

The aim of the study was to establish and verify a predictive vegetation model for plant community distribution in the alti-Mediterranean zone of the Lefka Ori massif, western Crete. Based on previous work three variables were identified as significant determinants of plant community distribution, namely altitude, slope angle and geomorphic landform. The response of four community types against these variables was tested using classification trees analysis in order to model community type occurrence. V-fold cross-validation plots were used to determine the length of the best fitting tree. The final 9node tree selected, classified correctly 92.5% of the samples. The results were used to provide decision rules for the construction of a spatial model for each community type. The model was implemented within a Geographical Information System (GIS) to predict the distribution of each community type in the study site. The evaluation of the model in the field using an error matrix gave an overall accuracy of 71%. The user's accuracy was higher for the Crepis-Cirsium (100%) and Telephium-Herniaria community type (66.7%) and relatively lower for the Peucedanum-Alyssum and Dianthus-Lomelosia community types (63.2% and 62.5%, respectively). Misclassification and field validation points to the need for improved geomorphological mapping and suggests the presence of transitional communities between existing community types.

Plant species and functional group effects on abiotic and microbial soil properties and plant-soil feedback responses in two grasslands

1 Plant species differ in their capacity to influence soil organic matter, soil nutrient availability and the composition of soil microbial communities. Their influences on soil properties result in net positive or negative feedback effects, which influence plant performance and plant community composition. 2 For two grassland systems, one on a sandy soil in the Netherlands and one on a chalk soil in the United Kingdom, we investigated how individual plant species grown in monocultures changed abiotic and biotic soil conditions. Then, we determined feedback effects of these soils to plants of the same or different species. Feedback effects were analysed at the level of plant species and plant taxonomic groups (grasses vs. forbs). 3 In the sandy soils, plant species differed in their effects on soil chemical properties, in particular potassium levels, but PLFA (phospholipid fatty acid) signatures of the soil microbial community did not differ between plant species. The effects of soil chemical properties were even greater when grasses and forbs were compared, especially because potassium levels were lower in grass monocultures. 4 In the chalk soil, there were no effects of plant species on soil chemical properties, but PLFA profiles differed significantly between soils from different monocultures. PLFA profiles differed between species, rather than between grasses and forbs. 5 In the feedback experiment, all plant species in sandy soils grew less vigorously in soils conditioned by grasses than in soils conditioned by forbs. These effects correlated significantly with soil chemical properties. None of the seven plant species showed significant differences between performance in soil conditioned by the same vs. other plant species. 6 In the chalk soil, Sanguisorba minor and in particular Briza media performed best in soil collected from conspecifics, while Bromus erectus performed best in soil from heterospecifics. There was no distinctive pattern between soils collected from forb and grass monocultures, and plant performance could not be related to soil chemical properties or PLFA signatures. 7 Our study shows that mechanisms of plant-soil feedback can depend on plant species, plant taxonomic (or functional) groups and site-specific differences in abiotic and biotic soil properties. Understanding how plant species can influence their rhizosphere, and how other plant species respond to these changes, will greatly enhance our understanding of the functioning and stability of ecosystems.

Detecting local adaptation in widespread grassland species - the importance of scale and local plant community

1 Adaptation of plant populations to local environments has been shown in many species but local adaptation is not always apparent and spatial scales of differentiation are not well known. In a reciprocal transplant experiment we tested whether: (i) three widespread grassland species are locally adapted at a European scale; (ii) detection of local adaptation depends on competition with the local plant community; and (iii) local differentiation between neighbouring populations from contrasting habitats can be stronger than differentiation at a European scale. 2 Seeds of Holcus lanatus, Lotus corniculatus and Plantago lanceolata from a Swiss, Czech and UK population were sown in a reciprocal transplant experiment at fields that exhibit environmental conditions similar to the source sites. Seedling emergence, survival, growth and reproduction were recorded for two consecutive years. 3 The effect of competition was tested by comparing individuals in weeded monocultures with plants sown together with species from the local grassland community. To compare large-scale vs. small-scale differentiation, a neighbouring population from a contrasting habitat (wet-dry contrast) was compared with the 'home' and 'foreign' populations. 4 In P. lanceolata and H. lanatus, a significant home-site advantage was detected in fitness-related traits, thus indicating local adaptation. In L. corniculatus, an overall superiority of one provenance was found. 5 The detection of local adaptation depended on competition with the local plant community. In the absence of competition the home-site advantage was underestimated in P. lanceolata and overestimated in H. lanatus. 6 A significant population differentiation between contrasting local habitats was found. In some traits, this small-scale was greater than large-scale differentiation between countries. 7 Our results indicate that local adaptation in real plant communities cannot necessarily be predicted from plants grown in weeded monocultures and that tests on the relationship between fitness and geographical distance have to account for habitat-dependent small-scale differentiation. Considering the strong small-scale differentiation, a local provenance from a different habitat may not be the best choice in ecological restoration if distant populations from a more similar habitat are available.

Species divergence and trait convergence in experimental plant community assembly

Despite decades of research, it remains controversial whether ecological communities converge towards a common structure determined by environmental conditions irrespective of assembly history. Here, we show experimentally that the answer depends on the level of community organization considered. In a 9-year grassland experiment, we manipulated initial plant composition on abandoned arable land and subsequently allowed natural colonization. Initial compositional variation caused plant communities to remain divergent in species identities, even though these same communities converged strongly in species traits. This contrast between species divergence and trait convergence could not be explained by dispersal limitation or community neutrality alone. Our results show that the simultaneous operation of trait-based assembly rules and species-level priority effects drives community assembly, making it both deterministic and historically contingent, but at different levels of community organization.

Phyto-activity of biocides used to manipulate herbivory: tests of three pesticides on fourteen plant species

1. We tested three pesticides used for field manipulations of herbivory for direct phytoactive effects on the germination and growth of 14 herbaceous plant species selected to provide a range of life-history strategies and functional groups. 2. We report three companion experiments: (A) Two insecticides, chlorpyrifos (granular soil insecticide) and dimethoate (foliar spray), were applied in fully-factorial combination to pot-germinated individuals of 12 species. (B) The same fully-factorial design was used to test for direct effects on the germination of four herbaceous legumes. (C) The molluscicide, metaldehyde, was tested for direct effects on the germination and growth of six plant species. 3. The insecticides had few significant effects on growth and germination. Dimethoate acted only on growth stimulating Anisantha sterilis, Sonchus asper and Stellaria graminea. In contrast, chlorpyrifos acted on germination increasing the germination of Trifolium dubium and Trifolium pratense. There was also a significant interactive effect of chlorpyrifos and dimethoate on the germination of T pratense. However, all. effects were relatively small in magnitude and explanatory power. The molluscicide had no significant effect on plant germination or growth. 4. The small number and size of direct effects of the pesticides on plant performance is encouraging for the use of these pesticides in manipulative experiments on herbivory, especially for the molluscicide. However, a smatt number of direct (positive) effects of the insecticides on some plant species need to be taken into account when interpreting field manipulations of herbivory with these compounds, and emphasises the importance of conducting tests for direct phyto-active effects. (C) 2004 Elsevier GmbH. All rights reserved.

Climate vs. soil factors in local adaptation of two common plant species

Evolutionary theory suggests that divergent natural selection in heterogeneous environments can result in locally adapted plant genotypes. To understand local adaptation it is important to study the ecological factors responsible for divergent selection. At a continental scale, variation in climate can be important while at a local scale soil properties could also play a role. We designed an experiment aimed to disentangle the role of climate and ( abiotic and biotic) soil properties in local adaptation of two common plant species. A grass (Holcus lanatus) and a legume ( Lotus corniculatus), as well as their local soils, were reciprocally transplanted between three sites across an Atlantic-Continental gradient in Europe and grown in common gardens in either their home soil or foreign soils. Growth and reproductive traits were measured over two growing seasons. In both species, we found significant environmental and genetic effects on most of the growth and reproductive traits and a significant interaction between the two environmental effects of soil and climate. The grass species showed significant home site advantage in most of the fitness components, which indicated adaptation to climate. We found no indication that the grass was adapted to local soil conditions. The legume showed a significant home soil advantage for number of fruits only and thus a weak indication of adaptation to soil and no adaptation to climate. Our results show that the importance of climate and soil factors as drivers of local adaptation is species-dependent. This could be related to differences in interactions between plant species and soil biota.

Changing precipitation patterns alter plant community dynamics and succession in an ex-arable grassland

1. Changes in the frequency of extreme events, such as droughts, may be one of the most significant impacts of climate change for ecosystems. Models predict more frequent summer droughts in much of England: this paper investigates the impact on different types of plants in an ex-arable grassland community. 2. A long-term experiment simulated increased and decreased summer precipitation. Substantial interannual variation allowed the effects of summer drought to be tested in combination with wet and dry weather in other seasons. This is important, as climate models predict increased winter precipitation. 3. Total cover abundance in early summer increased with increasing water supply in the previous summer; there was no effect of winter precipitation. Productivity is therefore likely to decrease with more frequent summer droughts, with no mitigating effect of wetter winters. 4. The percentage cover of perennial grasses declined during a natural drought in 1995-97; this was exacerbated by the experimental drought treatment and reduced by supplemented rainfall. Simultaneously, short-lived ruderal species increased; this was greatest in drought treatments and least with supplemented rainfall. 4. These trends were subsequently reversed during several years of unusually wet weather, with perennial grasses increasing and short-lived forbs decreasing. This occurred even in experimentally droughted plots, and we propose that it resulted from rapid coverage of gaps during wet autumns and winters. 6. Deep-rooted species generally proved to be more drought resistant, but there were exceptions. 7. We conclude that increased frequency of summer droughts could have serious implications for the establishment and successional development of ex-arable grasslands. Increased winter precipitation would moderate the impact on species composition, but not on productivity.

Conservation ecology of bees: populations, species and communities

Recent concerns regarding the decline of plant and pollinator species, and the impact on ecosystem functioning, has focused attention on the local and global threats to bee diversity. As evidence for bee declines is now accumulating from over broad taxonomic and geographic scales, we review the role of ecology in bee conservation at the levels of species, populations and communities. Bee populations and communities are typified by considerable spatiotemporal variation; whereby autecological traits, population size and growth rate, and plant-pollinator network architecture all play a role in their vulnerability to extinction. As contemporary insect conservation management is broadly based on species- and habitat-targeted approaches, ecological data will be central to integrating management strategies into a broader, landscape scale of dynamic, interconnected habitats capable of delivering bee conservation in the context of global environmental change.

Aggregation of parasitism risk in an aphid-parasitoid system: Effects of plant patch size and aphid density

Few studies have linked density dependence of parasitism and the tritrophic environment within which a parasitoid forages. In the non-crop plant-aphid, Centaurea nigra-Uroleucon jaceae system, mixed patterns of density-dependent parasitism by the parasitoids Aphidius funebris and Trioxys centaureae were observed in a survey of a natural population. Breakdown of density-dependent parasitism revealed that density dependence was inverse in smaller colonies but direct in large colonies (>20 aphids), suggesting there is a threshold effect in parasitoid response to aphid density. The CV2 of searching parasitoids was estimated from parasitism data using a hierarchical generalized linear model, and CV2>1 for A. funebris between plant patches, while for T. centaureae CV2>1 within plant patches. In both cases, density independent heterogeneity was more important than density-dependent heterogeneity in parasitism. Parasitism by T. centaureae increased with increasing plant patch size. Manipulation of aphid colony size and plant patch size revealed that parasitism by A. funebris was directly density dependent at the range of colony sizes tested (50-200 initial aphids), and had a strong positive relationship with plant patch size. The effects of plant patch size detected for both species indicate that the tritrophic environment provides a source of host density independent heterogeneity in parasitism, and can modify density-dependent responses. (c) 2007 Gessellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Phylogeny and the hierarchical organization of plant diversity

R. H. Whittaker's idea that plant diversity can be divided into a hierarchy of spatial components from alpha at the within-habitat scale through beta for the turnover of species between habitats to gamma along regional gradients implies the underlying existence of alpha, beta, and gamma niches. We explore the hypothesis that the evolution of a, (3, and gamma niches is also hierarchical, with traits that define the a niche being labile, while those defining a and 7 niches are conservative. At the a level we find support for the hypothesis in the lack of close significant phylogenetic relationship between meadow species that have similar a niches. In a second test, a niche overlap based on a variety of traits is compared between congeners and noncongeners in several communities; here, too, there is no evidence of a correlation between a niche and phylogeny. To test whether beta and gamma niches evolve conservatively, we reconstructed the evolution of relevant traits on evolutionary trees for 14 different clades. Tests against null models revealed a number of instances, including some in island radiations, in which habitat (beta niche) and elevational maximum (an aspect of the gamma niche) showed evolutionary conservatism.

Social bees and food plant associations in the Nilgiri Biosphere Reserve, India

The diversity of social bees was assessed at 15 sites across five locations of the Nilgiri Biosphere Reserve, Western Ghats, India, from January to December 2007. We also conducted floristic analyses of local vegetation in each site using one-hectare sample plots. All woody species with a dbh (diameter at breast height) : 30 cm were recorded within the plots. A total area of 9.72 ha was assessed for floristic composition. Similarity of floristic composition between sites was determined using the Jaccard's distance measure and a dendrogram constructed based on the hierarchical clustering of floristic dissimilarities between sites. A Bee Importance Index (BII) was developed to give a measure of the bee diversity at each site. This index was a sum of the species richness of bee species in a site and their visitation frequencies to flowers, calculated as mean flower visits hour 1 within 2 focal patches within one hectare plots. The visits of bee species to flowers were also recorded. The Jaccard distance measure indicated that the montane sites were quite dissimilar to the low elevation sites in floristic diversity. The BII was 7-9 for the wet forest sites and ranged from 4-6 for drier forest sites. Seventy three plant species were identified as social bee plants and of them 45% were visited by one species of bee, 37% by two bee species and 18% by more than two bee species, indicating a certain degree of floral specialization among bees.

Phylogeny, biogeography, and ecology of Ficus section Malvanthera (Moraceae)

We conducted the first molecular phylogenetic study of Ficus section Malvanthera (Moraceae; subgenus Urostigma) based on 32 Malvanthera accessions and seven outgroups representing other sections of Ficus subgenus Urostigma. We used DNA sequences from the nuclear ribosomal internal and external transcribed spacers (ITS and ETS), and the glyceraldehyde-3-phosphate dehydrogenase (G3pdh) region. Phylogenetic analysis using maximum parsimony, maximum likelihood and Bayesian methods recovered a monophyletic section Malvanthera to the exclusion of the rubber fig, Ficus elastica. The results of the phylogenetic analyses do not conform to any previously proposed taxonomic subdivision of the section and characters used for previous classification are homoplasious. Geographic distribution, however, is highly conserved and Melanesian Malvanthera are monophyletic. A new subdivision of section Malvanthera reflecting phylogenetic relationships is presented. Section Malvanthera likely diversified during a period of isolation in Australia and subsequently colonized New Guinea. Two Australian series are consistent with a pattern of dispersal out of rainforest habitat into drier habitats accompanied by a reduction in plant height during the transition from hemi-epiphytic trees to lithophytic trees and shrubs. In contradiction with a previous study of Pleistodontes phylogeny suggesting multiple changes in pollination behaviour, reconstruction of changes in pollination behaviour on Malvanthera, suggests only one or a few gains of active pollination within the section. (C) 2008 Elsevier Inc. All rights reserved.