Zaal Kikvidze

Alpine cushion plants inhibit the loss of phylogenetic diversity in severe environments

Biotic interactions can shape phylogenetic community structure (PCS). However, we do not know how the asymmetric effects of foundation species on communities extend to effects on PCS. We assessed PCS of alpine plant communities around the world, both within cushion plant foundation species and adjacent open ground, and compared the effects of foundation species and climate on alpha (within-microsite), beta (between open and cushion) and gamma (open and cushion combined) PCS. In the open, alpha PCS shifted from highly related to distantly related with increasing potential productivity. However, we found no relationship between gamma PCS and climate, due to divergence in phylogenetic composition between cushion and open sub-communities in severe environments, as demonstrated by increasing phylo-beta diversity. Thus, foundation species functioned as micro-refugia by facilitating less stress-tolerant lineages in severe environments, erasing a global productivity - phylogenetic diversity relationship that would go undetected without accounting for this important biotic interaction.

Facilitation in subnival vegetation patches

. We examined spatial relationships among species in the subnival zone of the central Caucasus. The species composition of 300 vegetation patches was analysed. 144 of them contained only one species, whereas the other 156 contained 2.36 ± 1.31 species, with species numbers distributed as follows: 59 patches with two species, 41 with three, 39 with four and 17 with five species. In the multi-species patches, the 22 most frequent species were examined and 46 statistically significant species associations, 36 positive and 10 negative, were found. Ten of these 22 species were typical subnival plants very rarely occurring at lower altitudes. The other 12 species are ‘invaders’ as they have broader altitudinal ranges of occurrence and are common in alpine and even in subalpine belts. Contrary to the typical subnival species the invaders were found exclusively in the patches with more than one species. Invader species were significantly more associated with subnival species while they were significantly less associated with other species than expected by chance. Our interpretation is that typical subnival species nurse plants from lower altitudes and facilitate their invasion to more adverse subnival environments.

Plant species associations in alpine‐subnival vegetation patches in the Central Caucasus

. Patches of herbaceous vegetation found at the boundary of the alpine and subnival belts in the Central Caucasus (3000 m) were analysed for species composition, interspecific associations, as well as for relations between age, size and diversity of the patches. 34 plant species were recorded. Positive and negative associations were found among the most frequent species. A model is presented that describes the cumulation of species in patches during succession. In the final and stable successional stage, the maximum number of species within a patch was nine. Species composition varies in patches of different age, typical alpine species being more frequent in older ones. The results are interpreted as evidence for niche differentiation and selection of species composition through the sieve’ of interspecific relations, taking place during succession in the studied patches.

Ecological Facilitation May Drive Major Evolutionary Transitions

There is a growing consensus among ecologists that ecological facilitation comprises a historically overlooked but crucial suite of biotic interactions. Awareness of such positive interactions has recently led to substantial modifications in ecological theory. In this article we suggest how facilitation may be included in evolutionary theory. Natural selection based on competition provides a conceptually complete paradigm for speciation, but not for major evolutionary transitions—the emergence of new and more complex biological structures such as cells, organisms, and eusocial populations. We find that the successful theories developed to solve these specific problematic transitions show a consistent pattern: they focus on positive interactions. We argue that facilitation between individuals at different levels of biological organization can act as a cohesive force that generates a new level of organization with higher complexity and thus allows for major evolutionary transitions at all levels of biological hierarchy.

The context dependence of beneficiary feedback effects on benefactors in plant facilitation

Facilitative effects of some species on others are a major driver of biodiversity. These positive effects of a benefactor on its beneficiary can result in negative feedback effects of the beneficiary on the benefactor and reduced fitness of the benefactor. However, in contrast to the wealth of studies on facilitative effects in different environments, we know little about whether the feedback effects show predictable patterns of context dependence. We reanalyzed a global data set on alpine cushion plants, previously used to assess their positive effects on biodiversity and the nature of the beneficiary feedback effects, to specifically assess the context dependence of how small- and large-scale drivers alter the feedback effects of cushion-associated (beneficiary) species on their cushion benefactors using structural equation modelling. The effect of beneficiaries on cushions became negative when beneficiary diversity increased and facilitation was more intense. Local-scale biotic and climatic conditions mediated these community-scale processes, having indirect effects on the feedback effect. High-productivity sites demonstrated weaker negative feedback effects of beneficiaries on the benefactor. Our results indicate a limited impact of the beneficiary feedback effects on benefactor cushions, but strong context dependence. This context dependence may help to explain the ecological and evolutionary persistence of this widespread facilitative system.

Facilitation among plants as an insurance policy for diversity in Alpine communities

Summary Positive interactions have started to gain a place as important drivers of community structure and biological diversity. Defined as non-trophic interspecific interactions that increase the average individual fitness of one species, by definition, the presence of one plant species enhances the chances that another species co-occur in the same place, indicating that positive interactions may determine biological diversity. However, this has been poorly explored. The majority of the studies addressing community-level consequences of facilitation have compared the diversity of the plant assemblages growing within nurses vs. those growing outside them, reporting contrasting results among them. Nonetheless, nurses and their alternative microhabitats (open areas among nurses) are part of the same community. Thus, if nurses allow for the persistence of species that otherwise would be excluded from the community, a net increase in the species diversity at the entire community level will be generated even though nurse plants contained fewer species than open areas. Here, we conducted a bibliographic search using the ISI Web of Knowledge data base and reviewed the literature conducted on alpine plant communities where assessments of the diversity of plants growing within and outside a nurse species were available. In most cases nurse species substantially increased species richness at the community level, despite the fact that in some cases, they contained lower species numbers than open areas. Nurse species enhanced species richness more in systems with impoverished local diversity, suggesting that facilitative interactions in alpine habitats act as an insurance policy that sustains diversity under very harsh conditions.

Combining observational and experimental methods in plant–plant interaction research

Background: Neighbour-removal experiments (NRE) and spatial pattern analyses (SPA) are commonly used methods to investigate plant–plant interactions. Although they address the same issue, they measure different aspects of plant interactions: experiments indicate contemporary processes, whilst observations of spatial patterns integrate the results of interactions that have prevailed in the past. Aim: The aim of this study was to propose a new conceptual approach that takes into account the chronological order between processes and the arising patterns i.e., the time lag between neighbour effects quantified with NRE and SPA, to detect shifts in the balance of plant interactions due to current environmental change. Methods: This conceptual approach was applied to alpine snowbeds. Data from NRE were used to calculate the importance index of current neighbour interactions. Spatial patterns were quantified using variance ratio statistics and were assumed to reflect historic interactions. Results: The results of the two approaches showed a consistent difference in the prevailing type of plant interactions and suggested a shift towards competition in recent times. Conclusions: The simultaneous application of NRE and SPA allows the detection of a recent shift in the balance of plant interactions and provides a deeper and more accurate insight into the temporal dynamics of plant communities that could not be gained using one method alone.

Loss of traditional knowledge aggravates wolf–human conflict in Georgia (Caucasus) in the wake of socio-economic change

Reports of the damage from wolf attacks have increased considerably over the last decade in Georgia (in the Caucasus). We interviewed locals about this problem in two focal regions: the Lanchkhuti area (in western Georgia) and Kazbegi District (in eastern Georgia) where livestock numbers had increased by an order of magnitude owing to dramatic shifts in the local economies over the last decade. This coincided with expanding habitats for wolves (abandoned plantations, for example). We found that the perceived damage from wolves was positively correlated with a poor knowledge of wolf habits and inappropriate livestock husbandry practices. Our results suggest a loss of traditional knowledge contributes strongly to the wolf–human conflicts in Georgia. Restoring traditional, simple but good practices—such as protecting herds using shepherd dogs and introducing bulls into the herds—can help one solve this problem.

Removal of Perennial Herbaceous Species Affects Response of Cold Desert Shrublands to Fire

Questions Two of the primary global change factors that threaten shrublands worldwide are loss of native perennial herbaceous species due to inappropriate livestock grazing and loss of native shrubs due to altered fire regimes. We asked: (1) How do the separate and interacting effects of removal of perennial herbaceous species and burning influence relative abundance of plant functional groups over longer time frames? (2) How do interactions between perennial herbaceous species removal and burning differ along environmental gradients? We discuss implications of our findings for ecosystem resilience to these disturbances. Location Shoshone Mountain Range, Nevada and East Tintic Range, Utah, western USA. Methods We used a factorial experiment to test effects of perennial herbaceous species removal (0%, 50%, and 100%) and burning (burned and not burned) on plant functional group cover along elevation gradients within watersheds characterized by Artemisia tridentata Nutt. vegetation types. The experiment was conducted in two locations (Nevada and Utah) with sites located at low (1960 and 1710 m), mid (2190 and 2085 m), and high (2380 and 2274 m) elevations and was repeated in two years. Percentage cover of native and exotic species and canopy area, density, and size of the shrub, A. tridentata, were evaluated 12 and 13 yrs after study implementation. Results Over a decade later, removal treatments resulted in highly significant decreases (40 to 62%) in perennial native grass and forb cover across site elevations. Burning decreased overall shrub and A. tridentata cover, but effects on perennial native grass cover differed among elevations. Removal had strong positive effects on A. tridentata seedling recruitment and resulted in progressive increases in density and canopy area following burning. A. tridentata canopy areas on burned plots with 0%, 50%, and 100% removal were 0.19, 0.40, and 0.84 m2, respectively. Annual invasive grass density also increased with degree of removal, and both density and cover decreased with elevation. Conclusions Our results show that loss of perennial herbaceous species, which can result from inappropriate livestock grazing, and loss of shrubs, which often results from fire, interact to affect key functional groups. The implications are that ecosystem resilience to disturbance in Cold Desert shrublands decreases when competition from perennial native grasses and forbs for available resources no longer prevents dominance by A. tridentata and other shrubs and/or annual invasive grasses. Managing livestock grazing to maintain or increase perennial herbaceous species, especially deep-rooted grasses which contribute to resilience along elevation gradients, can help prevent threshold crossings to undesirable states and retain critical ecosystem services following disturbances such as wildfire. This article is protected by copyright. All rights reserved.

A trait-based approach to understand the consequences of specific plant interactions for community structure

Question In plant communities, the presence of a species has consequences for other species, with some being competitively excluded while others benefit from the close vicinity of neighbours. Even though such specificity in plant interactions is common and known, there is no empirical assessment of the mechanisms that would help us understand its importance for plant diversity. Here we asked whether analysing spatial associations between plant traits known to affect the environment (i.e. effect traits) and those known to respond to the environment (i.e. response traits) might explain plant–plant interactions and their role in community assembly. Location Sierra Nevada Mountains, Spain Methods In a field study, we addressed the specificity of plant–plant interactions by quantifying effect traits of three co-occurring cushion-forming species and response traits of their associated plant assemblages. Traits were measured at the individual level and then aggregated to trait metrics (mean, range, dispersion) at the plot level. Finally, plot-level metrics of effect traits were related to response traits and the species composition of plant communities. Results Each cushion-forming species had a distinctive combination of effect traits and harboured a unique plant community with an exclusive composition of response traits. With multivariate statistics we showed that differences in effect traits (branch density and canopy height) among and within cushion species significantly explained response traits (specific leaf area, leaf dry matter content and plant height) of associated species and the local-scale species composition. Conclusions Using effect and response traits measured at the individual level, we provide a mechanistic understanding of the species-specificity of plant interactions and demonstrate how important such specificity is for species diversity in an ecosystem. This article is protected by copyright. All rights reserved.