Antarctic macrolichen modifies microclimate and facilitates vascular plants in the maritime Antarctica – a reply to Casanova‐Katny et al. (2014)

In a current article in the Journal of Vegetation Science, Casanova‐Katny et al. addressed a comment about an article by Molina‐Montenegro et al., which demonstrated the climate modification induced by the macrolichen Usnea antarctica and its role as facilitator. They provided useful corrections concerning species identification and pointed out several issues that, in their view, weakened our study. They indicated that the role of U. antarctica as a facilitative species in the maritime Antarctica is merely philosophical and has no ecological relevance. In this commentary, we argue why these critiques are unsubstantial, and provide evidence that the macrolichen can modify the microclimate, ameliorating the harsh conditions prevailing in Antarctica, establishing positive interactions and eventually facilitating vascular species. Thus, the macrolichen U. antarctica would act as a ‘nurse species’, playing a key role in structuring the maritime Antarctic plant community.     

Density may alter diversity–productivity relationships in experimental plant communities

Contemporary biodiversity experiments, in which plant species richness is manipulated and aboveground productivity of the system measured, generally demonstrate that lowering plant species richness reduces productivity. However, we propose that community density may in part compensate for this reduction of productivity at low diversity. We conducted a factorial experiment in which plant functional group richness was held constant at three, while plant species richness increased from three to six to 12 species and community density from 440 to 1050 to 2525 seedlings m⁻². Response variables included density, evenness and above- and belowground biomass at harvest. The density gradient converged slightly during the course of the experiment due to about 10% mortality at the highest sowing density. Evenness measured in terms of aboveground biomass at harvest significantly declined with density, but the effect was weak. Overall, aboveground, belowground and total biomass increased significantly with species richness and community density. However, a significant interaction between species richness and community density occurred for both total and aboveground biomass, indicating that the diversity–productivity relationship was flatter at higher than at lower density. Thus, high species richness enabled low-density communities to reach productivity levels otherwise seen only at high density. The relative contributions of the three functional groups C₃, C₄ and nitrogen-fixers to aboveground biomass were less influenced by community density at high than at low species richness. We interpret the interaction effects between community density and species richness on community biomass by expanding findings about constant yield and size variation from monocultures to plant mixtures.     

Positive associations between the cushion plant Arenaria polytrichoides (Caryophyllaceae) and other alpine plant species increase with altitude in the Sino‐Himalayas

Question: Does the facilitative effect of cushion plants increase with elevation as a result of increases in environmental harshness? Does this hypothesis apply in the Sino‐Himalayan Mountains? Location: Lakaka Pass on the Baima Snow Mountains (28°20′N, 99°05′E), SW China. Methods: We evaluated the spatial association of several plant species with the cushion plant Arenaria polytrichoides (Caryophyllaceae) at two elevations (4500 m and 4700 m) in the study site and monitored temperature, moisture and nutritional status of soil beneath and outside the cushions. Results: While 14 species grow more frequently associated with the cushions at the higher elevation, at the lower site only three species were positively associated with cushions. Eleven of the species that occurred at both elevations changed their spatial association from neutral or negative with cushions at the lower site to positive at the higher elevation site. Substrate temperatures were rather similar between the cushions and areas of bare ground. Cushions maintained higher moisture than areas of bare ground at both elevations. Soils beneath cushions contained significantly more available nitrogen and potassium compared to open areas at the higher elevation. Conclusions: Our results show that facilitation by A. polytrichoides cushions increases with elevation in the Sino‐Himalayan region. This facilitation effect of A. polytrichoides cushions is probably due to the improved nutrient availability provided by cushion plants in the higher elevation, and these conditions probably permit increased plant recruitment, growth and survival.     

Effects of plant diversity on invertebrate herbivory in experimental grassland

The rate at which a plant species is attacked by invertebrate herbivores has been hypothesized to depend on plant species richness, yet empirical evidence is scarce. Current theory predicts higher herbivore damage in monocultures than in species-rich mixtures. We quantified herbivore damage by insects and molluscs to plants in experimental plots established in 2002 from a species pool of 60 species of Central European Arrhenatherum grasslands. Plots differed in plant species richness (1, 2, 4, 8, 16, 60 species), number of functional groups (1, 2, 3, 4), functional group and species composition. We estimated herbivore damage by insects and molluscs at the level of transplanted plant individuals (“phytometer” species Plantago lanceolata, Trifolium pratense, Rumex acetosa) and of the entire plant community during 2003 and 2004. In contrast to previous studies, our design allows specific predictions about the relative contributions of functional diversity, plant functional identity, and species richness in relation to herbivory. Additionally, the phytometer approach is new to biodiversity-herbivory studies, allowing estimates of species-specific herbivory rates within the larger biodiversity-ecosystem functioning context. Herbivory in phytometers and experimental communities tended to increase with plant species richness and the number of plant functional groups, but the effects were rarely significant. Herbivory in phytometers was in some cases positively correlated with community biomass or leaf area index. The most important factor influencing invertebrate herbivory was the presence of particular plant functional groups. Legume (grass) presence strongly increased (decreased) herbivory at the community level. The opposite pattern was found for herbivory in T. pratense phytometers. We conclude that (1) plant species richness is much less important than previously thought and (2) plant functional identity is a much better predictor of invertebrate herbivory in temperate grassland ecosystems.     

Two approaches towards the relationship between plant species diversity and ecosystem functioning

Abstract. The main question to be dealt with in the papers published in this Special Feature is to which extent plant species richness can be applied as a parameter in restoration projects to qualify the ecosystem's state. Before considering this problem, it should be recognized that this approach illuminates only one side of the coin; the other side is touched by the opposite question, asking which plant species are essential components of an ecosystem. These two approaches towards the relationship between species richness and ecosystem functioning are not mutually exclusive, but should not be confused either. In view of ecosystem functioning certain species may be considered redundant, while in view of evolutionary processes certain ecosystem processes may be considered redundant. Where do the two approaches meet and when should they be separated? This paper touches upon this question by referring to the dual hierarchy of ecological systems.     

Improved water retention links high species richness with increased productivity in arctic tundra moss communities

A positive relationship between plant species richness and ecosystem functioning has been found in a number of experimental studies. Positive species interactions at high species numbers have been suggested as a cause, but mechanisms driving positive interactions have not often been tested. In this experiment we asked three questions: (1) What is the relationship between species richness and productivity in experimentally constructed moss communities? (2) Is this relationship affected by plant density? and (3) Can changes in moisture absorption and retention explain observed relationships? To answer these questions we exposed arctic tundra moss communities of different species richness levels (1–11 species) and two different densities in the greenhouse to two levels of drought (short and long). Biomass (by the community and individual species), height and community moisture absorption and retention were measured as response variables. High species diversity increased productivity (more so in low-density plots than in high-density plots), but only when plots were watered regularly. Plot moisture retention was improved at high species richness as well, and plant height and variation in height was increased compared to plants in monoculture. Under high-density and short-drought conditions 10 out of 12 species grew better in mixture than in monoculture, but under the long drought treatment only six species did. A positive feedback loop between biomass and improved humidity under high diversity was supported by path analysis. We conclude that in this community the relationship between species richness and productivity depends on moisture availability and density, with improved water absorption and retention likely to be the mechanism for increased plant growth when drought periods are short. Furthermore, since this is the opposite of what has been found for temperate moss communities, conclusions from one system cannot automatically be extrapolated to other systems.     

Ecosystem engineering affects ecosystem functioning in high-Andean landscapes

Ecosystem engineers are organisms that change the distribution of materials and energy in the abiotic environment, usually creating and maintaining new habitat patches in the landscape. Such changes in habitat conditions have been widely documented to affect the distributions and performances of other species but up to now no studies have addressed how such effects can impact the biotically driven physicochemical processes associated with these landscapes, or ecosystem functions. Based on the widely accepted positive relationship between species diversity and ecosystem functions, we propose that the effects of ecosystem engineers on other species could have an impact on ecosystem functions via two mutually inclusive mechanisms: (1) by adding new species into landscapes, hence increasing species diversity; and (2) by improving the performances of species already present in the landscape. To test these hypotheses, we focused on the effects of a high-Andean ecosystem engineer, the cushion plant Azorella monantha, by comparing the accumulation of plant biomass and nitrogen fixed in plant tissues as species richness increases in landscapes with and without the engineer species. Our results show that both ecosystem functions increased with species richness in both landscape types, but landscapes including A. monantha cushions reached higher outcomes of plant biomass and nitrogen fixed in plant tissues than landscapes without cushions. Moreover, our results indicate that such positive effects on ecosystem functions could be mediated by the two mechanisms proposed above. Then, given the conspicuousness of ecosystem engineering in nature and its strong influence on species diversity, and given the well-known relationship between species diversity and ecosystem function, we suggest that the application of the conceptual framework proposed herein to other ecosystems would help to advance our understanding of the forces driving ecosystem functioning. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Rapid decay of diversity-productivity relationships after invasion of experimental plant communities

So far, effects of species richness on ecosystem functioning have mainly been investigated in the short term in experimental communities from which invasion was prevented. We kept the local species pools of experimental grassland communities with 1, 2, 4, 8, and 32 species closed for five years and subsequently opened them for invasion by cessation of weeding. As long as communities were weeded, extinctions were rare but positively related to species richness, diversity-productivity relationships were positive, and more diverse systems had a greater temporal stability. Following cessation of weeding, species-poor communities were more prone to invasion. However, invasion increased extinction especially in species-rich communities. Within two years, differences in species richness and biomass production between sets of communities of different initial species richness disappeared and the positive diversity-productivity relationship was no longer detectable whereas species compositions remained distinct. This indicates that the positive diversity-productivity relationships during the weeding phase were mainly controlled by species richness.Bis anhin wurden die Effekte der Artenvielfalt auf das Funktionieren von Ökosystemen vor allem in kurzfristigen Experimenten untersucht, in denen die Einwanderung von Pflanzenarten in die bestehenden Gesellschaften verhindert wurde. Im vorliegenden Versuch wurden die lokalen Artenpools von 1, 2, 4, 8 und 32 Arten unserer experimentellen Graslandgesellschaften während 5 Jahren künstlich geschlossen gehalten und danach geöffnet indem nicht mehr gejätet wurde. Solange die Gesellschaften gejätet wurden, gab es wenige Aussterbeereignisse, die aber positiv mit der Artenvielfalt korreliert waren. Die Beziehung zwischen Diversität und Produktivität war positiv und Systeme höherer Diversität zeigten eine größere zeitliche Stabilität. Nach der Aufgabe des Jätens nahm die Einwanderung vor allem in artenarmen Gesellschaften zu. Die Einwanderung erhöhte jedoch besonders das Aussterben in ursprünglich artenreichen Gesellschaften. Innerhalb von zwei Jahren verschwanden die Unterschiede in der Artenzahl und Biomasseproduktion zwischen den verschiedenen Graslandgesellschaften und eine positive Beziehung zwischen Diversität und Produktivität war nicht mehr feststellbar. Die Artenzusammensetzung der Versuchsflächen blieb jedoch unterschiedlich. Das deutet darauf hin, daß die positive Beziehung zwischen Diversität und Produktivität während der ersten Phase des Experiments vor allem durch die Artenzahl und nicht durch die Artenzusammensetzung hervorgerufen wurde.     

Positive associations between the cushion plant Azorella monantha (Apiaceae) and alpine plant species in the Chilean Patagonian Andes

Low growing, compact cushion plants are a common and often dominant life form in temperate and subpolar alpine habitats. The cushion life-form can modify wind patterns, temperature and water availability and thus cushion species could be expected to act as nurse-plants facilitating the establishment of other alpine plant species on their surfaces. It has been suggested that the nurse effect should be most pronounced under more stressful environmental conditions, as found with increasing elevation in the alpine. One of the approaches used to detect the nurses has been the study of spatial associations among species, in which extreme clumping within or beneath one species has been interpreted as evidence of nursing. We characterized microclimatic conditions (soil and air temperature) within and outside cushions of Azorella monantha at two elevations (700 m a.s.l., corresponding to an elevation just above treeline, and 900 m a.s.l., corresponding to the upper limit of the cushion belt zone) on Cerro Diente in the Patagonian alpine of southern South America (50° S) and recorded all plant species growing upon cushions of various sizes and for paired sampling areas of equivalent sizes outside cushions. At 5 cm depth, soil temperature was slightly higher under cushions than under bare ground, but only significantly so at 900 m. Air temperature at ground level was significantly higher in the cushion microhabitat at both 700 m and 900 m, with the difference being more exaggerated at the highest elevation. At 700 m, a total of 27 species were recorded growing within cushions as compared to 29 outside cushions. At 900 m the corresponding numbers were 34 and 18. At the highest elevation, significantly more species grow within cushions than for equal areas outside cushions. Here moreover, 17 (48.6%) species grew preferentially within cushions, with eight of the latter being limited to the cushion microhabitat at this elevation. However, at 700 m there was no significant difference in species richness in the two microhabitats, and only one species (3.1%) grew preferentially on cushions. Considering individual species, nine occurring at both elevations showed non-preferential recruitment on cushions at 700 m, but significantly higher frequencies on cushions at 900 m. Results suggest striking altitudinal variation in the association with Azorella monantha on Cerro Diente, ranging from a very strong at 900 m to near absence at 700 m. Milder air and soil temperatures, shelter from wind, and greater water availability within cushions as opposed to outside cushions are discussed as possible factors favoring strong plant recruitment on cushions at higher elevations in the harsh Patagonian alpine environment. This revised version was published online in August 2006 with corrections to the Cover Date.     

Trait convergence and trait divergence in herbaceous plant communities: Mechanisms and consequences

In landscapes subject to intensive agriculture, both soil fertility and vegetation disturbance are capable of impacting strongly, evenly and simultaneously on the herbaceous plant cover and each tends to impose uniformity on the traits of constituent species. In more natural and ancient grasslands greater spatial and temporal variation in both productivity and disturbance occurs and both factors have been implicated in the maintenance of species‐richness in herbaceous communities. However, empirical data suggest that disturbance is the more potent driver of trait differentiation and species co‐existence at a local scale. This may arise from the great diversity in opportunities for establishment, growth or reproduction that arise when the intensity of competition is reduced by damage to the vegetation. In contrast to the diversifying effects of local disturbances, productivity‐related plant traits (growth rate, leaf longevity, leaf chemistry, leaf toughness, decomposition rate) appear to be less variable on a local scale. This difference in the effects of the productivity and disturbance filters arises from the relative constancy of productivity within the community and the diversity in agency and in spatial and temporal scales exhibited by disturbance events. Also, evolutionary responses to disturbances involve minor adaptive shifts in phenological and regenerative traits and are more likely to occur as micro‐evolutionary steps than the shifts in linked traits in the core physiology associated with the capacity to exploit productive and unproductive habitats. During the assembly of a community and over its subsequent lifespan filters with diversifying and convergent effects may operate simultaneously on recruitment from the local species pool and impose contrasted effects on the similarity of the trait values exhibited by co‐existing species. Moreover, as a consequence of the frequent association of productivity with the convergence filter, an additional difference is predicted in terms of the effects of the two filters on ecosystem functioning. Convergence in traits selected by the productivity filter will exert effects on both the plant community and the ecosystem while divergent effects of the disturbance filter will be restricted to the plant community.     

Additive and interactive effects of plant genotypic diversity on arthropod communities and plant fitness

Recent research suggests that genetic diversity in plant populations can shape the diversity and abundance of consumer communities. We tested this hypothesis in a field experiment by manipulating patches of Evening Primrose ( Oenothera biennis ) to contain one, four or eight plant genotypes. We then surveyed 92 species of naturally colonizing arthropods. Genetically diverse plant patches had 18% more arthropod species, and a greater abundance of omnivorous and predacious arthropods, but not herbivores, compared with monocultures. The effects of genotypic diversity on arthropod communities were due to a combination of interactive and additive effects among genotypes within genetically diverse patches. Greater genetic diversity also led to a selective feedback, as mean genotype fitness was 27% higher in diverse patches than in monocultures. A comparison between our results and the literature reveals that genetic diversity and species diversity can have similar qualitative and quantitative effects on arthropod communities. Our findings also illustrate the benefit of preserving genetic variation to conserve species diversity and interactions within multitrophic communities.     

Dominance–diversity relationships in ant communities differ with invasion

The relationship between levels of dominance and species richness is highly contentious, especially in ant communities. The dominance‐impoverishment rule states that high levels of dominance only occur in species‐poor communities, but there appear to be many cases of high levels of dominance in highly diverse communities. The extent to which dominant species limit local richness through competitive exclusion remains unclear, but such exclusion appears more apparent for non‐native rather than native dominant species. Here we perform the first global analysis of the relationship between behavioral dominance and species richness. We used data from 1,293 local assemblages of ground‐dwelling ants distributed across five continents to document the generality of the dominance‐impoverishment rule, and to identify the biotic and abiotic conditions under which it does and does not apply. We found that the behavioral dominance–diversity relationship varies greatly, and depends on whether dominant species are native or non‐native, whether dominance is considered as occurrence or relative abundance, and on variation in mean annual temperature. There were declines in diversity with increasing dominance in invaded communities, but diversity increased with increasing dominance in native communities. These patterns occur along the global temperature gradient. However, positive and negative relationships are strongest in the hottest sites. We also found that climate regulates the degree of behavioral dominance, but differently from how it shapes species richness. Our findings imply that, despite strong competitive interactions among ants, competitive exclusion is not a major driver of local richness in native ant communities. Although the dominance‐impoverishment rule applies to invaded communities, we propose an alternative dominance‐diversification rule for native communities.     

Effects of plant diversity,community composition and environmental parameters on productivity in montane European grasslands

In the past years, a number of studies have used experimental plant communities to test if biodiversity influences ecosystem functioning such as productivity. It has been argued, however, that the results achieved in experimental studies may have little predictive value for species loss in natural ecosystems. Studies in natural ecosystems have been equivocal, mainly because in natural ecosystems differences in diversity are often confounded with differences in land use history or abiotic parameters. In this study, we investigated the effect of plant diversity on ecosystem functioning in semi-natural grasslands. In an area of 10×20 km, we selected 78 sites and tested the effects of various measures of diversity and plant community composition on productivity. We separated the effects of plant diversity on ecosystem functioning from potentially confounding effects of community composition, management or environmental parameters, using multivariate statistical analyses. In the investigated grasslands, simple measures of biodiversity were insignificant predictors of productivity. However, plant community composition explained productivity very well (R²=0.31) and was a better predictor than environmental variables (soil and site characteristics) or management regime. Thus, complex measures such as community composition and structure are important drivers for ecosystem functions in semi-natural grasslands. Furthermore, our data show that it is difficult to extrapolate results from experimental studies to semi-natural ecosystems, although there is a need to investigate natural ecosystems to fully understand the relationship of biodiversity and ecosystem functioning.     

Microbial phylotype composition and diversity predicts plant productivity and plant–soil feedbacks

The relationship between ecological variation and microbial genetic composition is critical to understanding microbial influence on community and ecosystem function. In glasshouse trials using nine native legume species and 40 rhizobial strains, we find that bacterial rRNA phylotype accounts for 68% of amoung isolate variability in symbiotic effectiveness and 79% of host specificity in growth response. We also find that rhizobial phylotype diversity and composition of soils collected from a geographical breadth of sites explains the growth responses of two acacia species. Positive soil microbial feedback between the two acacia hosts was largely driven by changes in diversity of rhizobia. Greater rhizobial diversity accumulated in association with the less responsive host species, Acacia salicina, and negatively affected the growth of the more responsive Acacia stenophylla. Together, this work demonstrates correspondence of phylotype with microbial function, and demonstrates that the dynamics of rhizobia on host species can feed back on plant population performance.     

Nurse effect of the native cushion plant Azorella monantha on the invasive non-native Taraxacum officinale in the high-Andes of central Chile

Positive interactions among native plant species are common in alpine habitats, particularly those where one species (nurse plant) generates microclimatic conditions that are more benign than the surrounding environment, facilitating the establishment of other species. Nonetheless, these microclimatic conditions could facilitate the establishment of non-native species as well. A conspicuous component of the alien alpine flora of the central Chilean Andes is the perennial herb Taraxacum officinale agg. (dandelion). In contrast to other alien species that are restricted to human-disturbed sites, T. officinale is frequently observed growing within native plant communities dominated by cushion plants. In this study we evaluated if T. officinale is positively associated with the cushion plant Azorella monantha. Via seedling survival experiments and gas-exchange measurements we also assessed the patterns of facilitation between cushions and dandelions, and explore the potential mechanisms of invasion by dandelions. T. officinale grows spatially positively associated with cushions of A. monantha. Survival of seedlings, as well as their net-photosynthetic rates and stomatal conductance, were higher within cushions than in open areas away from them, suggesting that the microclimatic modifications generated by this native cushion facilitates the establishment and performance of a non-native invasive species. Our results, as well as other recent studies, highlight the role of native communities in facilitating rather than constraining non-native plant invasions, particularly in stressful habitats such as alpine environments.     

How community adaptation affects biodiversity–ecosystem functioning relationships

Evidence is growing that evolutionary dynamics can impact biodiversity–ecosystem functioning (BEF) relationships. However the nature of such impacts remains poorly understood. Here we use a modelling approach to compare random communities, with no trait evolutionary fine‐tuning, and co‐adapted communities, where traits have co‐evolved, in terms of emerging biodiversity–productivity, biodiversity–stability and biodiversity–invasion relationships. Community adaptation impacted most BEF relationships, sometimes inverting the slope of the relationship compared to random communities. Biodiversity–productivity relationships were generally less positive among co‐adapted communities, with reduced contribution of sampling effects. The effect of community‐adaptation, though modest regarding invasion resistance, was striking regarding invasion tolerance: co‐adapted communities could remain very tolerant to invasions even at high diversity. BEF relationships are thus contingent on the history of ecosystems and their degree of community adaptation. Short‐term experiments and observations following recent changes may not be safely extrapolated into the future, once eco‐evolutionary feedbacks have taken place.