Trait dynamics of Mediterranean xerophytic shrub communities growing on stabilised inland dunes respond to nutrient and aridity gradients

Background: Understanding the processes that determine community assembly and their dynamics is a central issue in ecology. The analysis of functional diversity can improve our understanding of these dynamics by identifying community assembly processes.

Aims: We studied the effect of environment–community covariations on both functional diversity and functional structure of xerophytic shrub communities for inferring the community assembly processes shaping this vegetation type.

Methods: Functional diversity was quantified using (1) community-weighted mean of the studied traits, (2) functional groups, defined using Ward’s hierarchical agglomerative clustering method and (3) Rao’s quadratic entropy. Relationships between functional diversity and environmental gradients were identified by Spearman correlations and modelled using generalised additive models.

Results: Variations in community composition and functional diversity correlated with soil nutrient availability and aridity. Increasing nutrient availability resulted in both greater average plant height and higher abundance of plants with green photosynthetic organ colour, whereas the abundance of nanophanerophytes increases with aridity.

Conclusions: The species composition and trait structure of the studied Mediterranean xerophytic shrub communities varies along nutrient and aridity gradients. This supports the importance of environmental filters for the local assembly and dynamics of these inland dune communities.     

Functional diversity and composition of bryophyte water-related traits in Azorean native vegetation

Background: Functional trait-based approaches link species diversity patterns to ecosystem functioning. In the context of global change, understanding these links is vital for developing holistic biodiversity management strategies. Bryophytes, important ecosystem components owing to their biogeochemical functions, have not been the focus of many functional studies.

Aims: This is the first assessment of bryophyte functional diversity in the Azores archipelago, aiming to uncover multivariate trait richness and composition patterns along the elevational gradient on Terceira Island.

Methods: Based on five water acquisition and retention traits of leafy liverworts and mosses, we calculated functional diversity metrics within and among six bryophyte communities sampled along a 1021-m elevational transect.

Results: Trait composition differed significantly between coastal and mountain communities. Mosses presented inrolled leaves and ornamented leaf cells at low elevation but not at high elevation. These patterns were associated with an uphill shift from drier and warmer conditions to a moister and cooler environment.

Conclusions: Future climatic changes might affect bryophyte functional diversity patterns in Terceira Island, particularly for mosses. These results can be directly compared with those obtained for other archipelagos where the same protocol has been applied, allowing a joint assessment of insular vegetation functional diversity patterns.     

Changes in plant taxonomic and functional diversity patterns following treeline advances in the South Urals

Background: Treeline ecotones represent environmental boundaries that fluctuate in space and time and thus induce changes in plant taxonomic and functional diversity.

Aims: To study changes through time in taxonomic and functional plant diversity patterns along the treeline ecotone.

Methods: In 2002, vegetation was sampled along a gradient from upper montane forest to the treeline–alpine transition in the South Ural Mountains, Russia. In 2014, vegetation was resampled and plant functional traits were collected. We studied spatial and temporal changes in plant species composition, functional composition and functional diversity.

Results: Species composition and diversity changed along the elevational gradient. The functional composition in height, leaf area, specific leaf area and leaf nitrogen content decreased with elevation, whereas functional composition of leaf carbon content increased. We found a temporal shift towards shorter plants with smaller leaves in treeline sites. Functional richness varied in several traits along the elevational gradient, while functional dispersion showed a trend towards increased functional dispersion in height, specific leaf area and leaf nitrogen in the treeline–tundra transition.

Conclusions: Tree encroachment across the treeline ecotone has resulted in a shift in plant species relative abundances and functional diversity, possibly affecting plant community assembly patterns.     

Effects of environmental heterogeneity on species diversity and composition of terrestrial bryophyte assemblages in tropical montane forests of southern Ecuador

Background: Most studies on tropical bryophytes deal with epiphytic species. This is the first ecological study of tropical forests that focuses specifically on terrestrial bryophytes.

Aim: To investigate the differences between slope and ridge environments in upper montane forests of southern Ecuador in terms of species diversity (richness, abundance), species composition and life forms of terrestrial bryophytes.

Methods: We used Non-metric Multidimensional Scaling (NMDS) to group bryophyte relevés by study location, habitat type and exposure class. Species indicator values were calculated and compared for different habitats.

Results: In total, 140 species were recorded, the majority being liverworts. NMDS analyses and Mantel correlations clearly separated between slope and ridge relevés, and between sunny and shaded microhabitats on ridges. Bryophyte life forms also showed different distribution patterns in slope and in ridge habitats. Mosses were more prominent in sunny than in shaded microhabitats.

Conclusions: Environmental differentiation between ridges and slopes, and small-scale variation in microclimatic conditions caused by differences in exposure, were stronger predictors of species richness and composition than geographical distance between study sites.     

Elevation,Topography, and Edge Effects Drive Functional Composition of Woody Plant Species in Tropical Montane Forests

Tropical montane forests comprise heterogeneous environments along natural gradients of topography and elevation. Human‐induced edge effects further increase the environmental heterogeneity in these forests. The simultaneous effects of natural and human‐induced gradients on the functional diversity of plant leaf traits are poorly understood. In a tropical montane forest in Bolivia, we studied environmental gradients associated with elevation (from 1900 m to 2500 m asl), topography (ridge and gorge), and edge effects (forest edge vs. forest interior), and their relationship with leaf traits and resource‐use strategies. First, we investigated associations of environmental conditions (soil properties and microclimate) with six leaf traits, measured on 119 woody plant species. Second, we evaluated changes in functional composition with community‐weighted means and functional structure with multidimensional functional diversity indices (FRic, FEve and FDiv). We found significant associations between leaf traits and soil properties in accordance with the trade‐off between acquisition and conservation of resources. Functional composition of leaf traits shifted from the dominance of acquisitive species in habitats at low altitudes, gorges, and forest interior to the dominance of conservative species in habitats at high altitudes, ridges, and forest edges. Functional structure was only weakly associated with the environmental gradients. Natural and human‐induced environmental gradients, especially soil properties, are important for driving leaf traits and resource‐use strategies of woody plants. Nevertheless, weak associations between functional structure and environmental gradients suggest a high redundancy of functional leaf traits in this tropical montane forest.     

Plant–environment interactions through a functional traits perspective: a review of Italian studies

Abstract

Italy is among the European countries with the greatest plant diversity due to both a great environmental heterogeneity and a long history of man–environment interactions. Trait-based approaches to ecological studies have developed greatly over recent decades worldwide, although several issues concerning the relationships between plant functional traits and the environment still lack sufficient empirical evaluation. To draw insights on the association between plant functional traits and direct and indirect human and natural pressures on the environmental drivers, this article summarizes the existing knowledge on this topic by reviewing the results of studies performed in Italy adopting a functional trait approach on vascular plants, bryophytes and lichens. Although we recorded trait measurements for 1418 taxa, our review highlighted some major gaps in plant traits knowledge: Mediterranean ecosystems are poorly represented; traits related to belowground organs are still overlooked; traits measurements for bryophytes and lichens are lacking. Finally, intraspecific variation has been little studied at community level so far. We conclude by highlighting the need for approaches evaluating trait–environment relationship at large spatial and temporal scales and the need of a more effective contribution to online databases to tie more firmly Italian researchers to international scientific networks on plant traits.     

Grassland plant species occurring in extensively managed road verges are filtered by urban environments

Background: Urbanisation filters species in communities depending on their adaptability to conditions in built-up areas, especially in semi-natural habitats. Roadside vegetation is widespread along urban-rural gradients and is therefore a good place to study landscape-scale factors influencing plant community composition.

Aim: Our study aimed to assess how plant species distributions vary between urban and rural landscape contexts and to identify biological traits favoured in urban areas.

Methods: Presence/absence data for 63 indigenous common species were collected in 296 road verge patches distributed along the urban–rural gradient in three French cities. We investigated the effects of landscape composition on species assemblages and related individual species responses to urbanisation to functional traits associated with dispersal and persistence capacity.

Results: Many grassland species were negatively affected by increasing proportion of built-up areas in the landscape. Insect-pollination and high seed production appeared to be key traits favoured in grassland communities in urban areas, whereas dispersal modes were less related to plant distribution.

Conclusions: This study has demonstrated that urban filters affect common species of widespread, managed road verges. Better knowledge of the flora of these herbaceous roadsides may contribute to the conservation of common biodiversity within other grassland habitats found in urban areas.     

Patterns and origin of intraspecific functional variability in a tropical alpine species along an altitudinal gradient

Background : Intraspecific functional variability (IFV) along altitudinal gradients is a powerful proxy to infer the responses of plants to abrupt environmental changes. We envisage that IFV shows distinctive patterns in tropical and extratropical alpine regions.

Aims : To characterise the patterns and explore the origin of IFV in a tropical alpine species in a context of upward range extension.

Methods : We examined variations in a series of plant functional traits in Lasiocephalus ovatus, inside and outside a nurse plant along a 600 m altitudinal gradient in the Ecuadorian Andes, and we studied its genetic variability.

Results : More conservative traits were developed at higher elevation, in contrast to extratropical alpine plants, which commonly develop opportunistic traits in response to late snowmelt close to their upper altitudinal limit. The presence of nurse cushions did not alter this trend. Increasing genetic distance along the gradient suggested that IFV might be partly genetically induced.

Conclusions : Our data combined with existing literature in tropical alpine environments lead the way to a stimulating scientific challenge: determining if patterns of plant altitudinal distribution in tropical alpine areas in response to climate change are predictable from patterns described in extratropical alpine areas.     

Plant Trait Variation along an Altitudinal Gradient in Mediterranean High Mountain Grasslands: Controlling the Species Turnover Effect

Assessing changes in plant functional traits along gradients is useful for understanding the assembly of communities and their response to global and local environmental drivers. However, these changes may reflect the effects of species composition (i.e. composition turnover), species abundance (i.e. species interaction), and intra-specific trait variability (i.e. species plasticity). In order to determine the relevance of the latter, trait variation can be assessed under minimal effects of composition turnover. Nine sampling sites were established along an altitudinal gradient in a Mediterranean high mountain grassland community with low composition turnover (Madrid, Spain; 1940 m–2419 m). Nine functional traits were also measured for ten individuals of around ten plant species at each site, for a total of eleven species across all sites. The relative importance of different sources of variability (within/between site and intra-/inter-specific functional diversity) and trait variation at species and community level along the considered gradients were explored. We found a weak individual species response to altitude and other environmental variables although in some cases, individuals were smaller and leaves were thicker at higher elevations. This lack of species response was most likely due to greater within- than between-site species variation. At the community level, inter-specific functional diversity was generally greater than the intra-specific component except for traits linked to leaf element content (leaf carbon content, leaf nitrogen content, δ¹³C and δ¹⁵N). Inter-specific functional diversity decreased with lower altitude for four leaf traits (specific leaf area, leaf dry matter content, δ¹³C and δ¹⁵N), suggesting trait convergence between species at lower elevations, where water shortage may have a stronger environmental filtering effect than colder temperatures at higher altitudes. Our results suggest that, within a vegetation type encompassing various environmental gradients, both, changes in species abundance and intra-specific trait variability adjust for the community functional response to environmental changes.     

Processes of community assembly in an environmentally heterogeneous,high biodiversity region

Despite decades of study, the relative importance of niche‐based versus neutral processes in community assembly remains largely ambiguous. Recent work suggests niche‐based processes are more easily detectable at coarser spatial scales, while neutrality dominates at finer scales. Analyses of functional traits with multi‐year multi‐site biodiversity inventories may provide deeper insights into assembly processes and the effects of spatial scale. We examined associations between community composition, species functional traits, and environmental conditions for plant communities in the Kouga‐Baviaanskloof region, an area within South Africa's Cape Floristic Region (CFR) containing high α and β diversity. This region contains strong climatic gradients and topographic heterogeneity, and is comprised of distinct vegetation classes with varying fire histories, making it an ideal location to assess the role of niche‐based environmental filtering on community composition by examining how traits vary with environment. We combined functional trait measurements for over 300 species with observations from vegetation surveys carried out in 1991/1992 and repeated in 2011/2012. We applied redundancy analysis, quantile regression, and null model tests to examine trends in species turnover and functional traits along environmental gradients in space and through time. Functional trait values were weakly associated with most spatial environmental gradients and only showed trends with respect to vegetation class and time since fire. However, survey plots showed greater compositional and functional stability through time than expected based on null models. Taken together, we found clear evidence for functional distinctions between vegetation classes, suggesting strong environmental filtering at this scale, most likely driven by fire dynamics. In contrast, there was little evidence of filtering effects along environmental gradients within vegetation classes, suggesting that assembly processes are largely neutral at this scale, likely the result of very high functional redundancy among species in the regional species pool.     

High plant taxonomic beta diversity and functional and phylogenetic convergence between two Neotropical inselbergs

ABSTRACT

Background: Inselbergs (granitic and gneissic rock outcrops) are common elements in the Atlantic Forest and present large taxonomic (TD), functional (FD) and phylogenetic (PD) diversity.

Aims: We investigated how plant diversity changed across ecological and biogeographic scales by comparing TD, FD and PD of communities within and between two inselbergs. We expected converging FD and PD but distinct TD between outcrops, because of similar local environmental conditions in inselbergs and the long-term lineage isolation.

Methods: We calculated TD, PD and FD, and partitioned diversity into α (each inselberg), β (between inselbergs) and γ (whole sample) components. Phylogenetic signal was estimated for all traits. To link environmental predictors to functional traits a redundancy analysis was run. Variation in TD, FD and PD was analysed by general linear models with patch area and the two inselbergs as predictors.

Results: The inselbergs were taxonomically different, but showed convergence in their functional and phylogenetic diversity. The limited retention of phylogenetic signal suggests that different species may converge and respond similarly to environmental variables. Within inselbergs, larger patches displayed higher TD, FD and PD.

Conclusions: Seeking conservation strategies for inselbergs is challenging since, despite their functional and phylogenetic similarity, endemic species make individual rock outcrops unique.     

Long-term vegetation stability in northern Europe as assessed by changes in species co-occurrences

Background: The effect of the anticipated climate change on the stability of vegetation and the factors underlying this stability are not well understood.

Aims: Our objective was to quantify long-term vegetation changes in a range of habitats in northern Europe by exploring species co-occurrences and their links to diversity and productivity gradients.

Methods: We re-sampled vegetation in 16 arctic, mountain and mire sites 20 to 90 years after the original inventories. A site-specific change in species assemblages (stability) was quantified using species co-occurrences. Using a randomisation test we tested whether the changes observed were significantly greater than those expected by chance. Relationships between patterns in vegetation stability and time between surveys, numbers of plots, or species diversity and proxies for productivity, were tested using regression analysis.

Results: At most sites the changes in species co-occurrences of vascular plants and bryophytes were greater than those expected by chance. The changes observed were found to be unrelated to gradients in productivity or diversity.

Conclusions: Changes in species co-occurrences are not strongly linked to diversity or productivity gradients in vegetation, suggesting that other gradients or site-specific factors (e.g. land use or species interactions) may be more important in controlling recent compositional shifts in vegetation in northern Europe.     

Notes On The Occurrence Of Clathrus Cancellatus,Tournf., In Argyllshire

Background: Estuaries are characterised by salinity gradients and regular flooding events. These environmental factors form stress gradients, along which species composition changes.

Aims: Analyse and compare patterns of plant species diversity along the estuarine salinity and flooding gradients of the Elbe and Connecticut Rivers.

Methods: Vegetation was sampled at three elevations (low, mid, high) in five sites of each marsh type (fresh, brackish, salt) in both estuaries. Patterns of species density (SD) and evenness (E) along the gradients were analysed and compared between the two estuaries with three-factor ANOVAs.

Results: The regional species pool was 33% higher for the Connecticut than for the Elbe. SD of fresh marshes (19 ± 2.2) was more than twice in the Connecticut than in the Elbe. We found an overall increase in SD from low to high elevation and from salt to freshwater marshes in both estuaries. However, SD and E were strongly depressed at intermediate elevations in the Elbe fresh and brackish marshes.

Conclusions: Although diversity patterns in the two estuaries show overall similarities, patterns of SD and E differ, when particular elevational zones and marsh types are compared. We hypothesise this to be due to evolutionary and historical influences on the regional species pools, shaping the impact of local biotic and abiotic processes.     

Functional traits reveal environmental constraints on amphibian community assembly in a subtropical dry forest

The relationships between functional traits and environmental gradients are useful to identify different community assembly processes. In this work, we used an approach based on functional traits to analyse if changes in hydroperiod and tree covers of ponds are relevant for local amphibian community assembly processes. Ephemeral ponds with low vegetation cover are expected to impose constraints on different species with particular trait combinations and, therefore, to exhibit communities with lower functional diversity than more stable ponds with greater tree cover. Sampling was conducted in 39 temporary ponds located along vegetation and hydroperiod gradients in the most arid portion of the Chaco ecoregion. Seven functional traits were measured in each species present in the regional pool. Associations between these traits and environmental gradients were detected using multivariate ordination techniques and permutation test (RLQ and fourth‐corner analyses respectively). Functional diversity indices were then calculated and related to variations in the environmental gradients. The results obtained allowed us to identify different sets of traits associated with hydroperiod and tree cover, suggesting that these environmental variables are relevant for structuring amphibian communities according to interspecific variations in functional traits from both, larval and adult stages. Contrary to our expectations, communities associated with more stable ponds and with greater tree cover exhibited lower functional diversity than expected by chance (and were the ponds with highest species richness). This result indicates that the reduction in relative importance of environmental restrictions imposed by a very short hydroperiod and the lack of tree cover, favours different species of the regional pool that are similar in several functional traits. Accordingly, communities associated with stable ponds with high tree cover exhibited high functional redundancy.     

Liverworts of southwest Ethiopian montane forests: ecological and biogeographical notes

Abstract

Ethiopia has diverse topographic features and climatic conditions with a diverse flora. The liverwort flora of southwest Ethiopia is practically unknown, despite a favourable climate and the occurrence of suitable ecosystems such as montane rainforests. During an ecological study of diversity patterns of bryophytes and vascular plants in relation to land use, we recorded many bryophyte species. In this paper we report the finding of 89 species of liverworts, and give short ecological notes and describe the distribution (locally and in Africa) for each species. Of these, 51 species are newly reported from Ethiopia. It is thus obvious that Ethiopia is considerably richer in liverworts than might be expected from previous checklists.     

Phytohormone Profiling across the Bryophytes

BackgroundBryophytes represent a very diverse group of non-vascular plants such as mosses, liverworts and hornworts and the oldest extant lineage of land plants. Determination of endogenous phytohormone profiles in bryophytes can provide substantial information about early land plant evolution. In this study, we screened thirty bryophyte species including six liverworts and twenty-four mosses for their phytohormone profiles in order to relate the hormonome with phylogeny in the plant kingdom.MethodologySamples belonging to nine orders (Pelliales, Jungermanniales, Porellales, Sphagnales, Tetraphidales, Polytrichales, Dicranales, Bryales, Hypnales) were collected in Central and Northern Bohemia. The phytohormone content was analysed with a high performance liquid chromatography electrospray tandem-mass spectrometry (HPLC-ESI-MS/MS).ConclusionThe apparent differences in conjugation and/or degradation strategies of growth hormones between liverworts and mosses might potentially show a hidden link between vascular plants and liverworts. On the other hand, the complement of stress hormones in bryophytes probably correlate rather with prevailing environmental conditions and plant survival strategy than with plant evolution.     

Taxonomical vs. functional responses of bee communities to fire in two contrasting climatic regions

1. Valuable insights into mechanisms of community responses to environmental change can be gained by analysing in tandem the variation in functional and taxonomic composition along environmental gradients. 2. We assess the changes in species and functional trait composition (i.e. dominant traits and functional diversity) of diverse bee communities in contrasting fire-driven systems in two climatic regions: Mediterranean (scrub habitats in Israel) and temperate (chestnut forests in southern Switzerland). 3. In both climatic regions, there were shifts in species diversity and composition related to post-fire age. In the temperate region, functional composition responded markedly to fire; however, in the Mediterranean, the taxonomic response to fire was not matched by functional replacement. 4. These results suggest that greater functional stability to fire in the Mediterranean is achieved by replacement of functionally similar species (i.e. functional redundancy) which dominate under different environmental conditions in the heterogeneous landscapes of the region. In contrast, the greater functional response in the temperate region was attributed to a more rapid post-fire vegetation recovery and shorter time-window when favourable habitat was available relative to the Mediterranean. 5. Bee traits can be used to predict the functional responses of bee communities to environmental changes in habitats of conservation importance in different regions with distinct disturbance regimes. However, predictions cannot be generalized from one climatic region to another where distinct habitat configurations occur.     

Plant functional identity as the predictor of carbon storage in semi-arid ecosystems

Background: There is an increasing consensus that ecosystem processes are governed by functional identity and trait variation rather than species richness. Despite its importance, the relative effect of relevant functional traits for carbon storage has remained mostly untested in different bioclimatic regions.

Aims: In this study, different components of functional diversity such as community-weighted means of trait values (CWM), functional trait diversity (Rao’s quadratic diversity), functional richness (FRi), functional evenness (FEv) and functional divergence (FDiv) were used to associate carbon content of above-ground biomass, litter and soil in four bioclimatic regions including warm and cold-steppe, semi-steppe rangelands and oak dry forest in the south-west of Iran.

Methods: Several key important traits highly associated with carbon storage including specific leaf area (SLA), height (H), leaf dry matter content, leaf nitrogen and phosphorus content (LNC and LPC), leaf longevity, wood specific gravity and life form were determined to quantify single and multiple traits that contribute to different components of plant functional diversity.

Results: The results showed that CWM of H, Chamaephyte life form, LNC and LPC were among the most important aspects of functional diversity that positively predicted carbon storage in above-ground biomass and soil. We also observed the negative association of carbon storage with FEv of LNC, Rao of LNC and FEv of multiple traits in the rangelands and the negative association of carbon storage with FDiv of SLA in the forest.

Conclusions: Our results indicate that different components of functional diversity are essential for a mechanistic understanding of the role of plant diversity for carbon storage. The negative associations between FDiv and FEv and carbon storage do not provide support for the complementarity niche hypothesis. Our results suggest that in the more functionally diverse ecosystems dominated by functionally important species with key traits, the so-called functional identity does indeed promote carbon storage, at least in these semi-arid ecosystems.     

Plant functional strategies and environmental constraints in Mediterranean high mountain grasslands in central Spain

Background: The relationship between plant traits and environmental factors will be of value in understanding of functional strategies that plants have developed to cope with the environmental constraints on plant life in Mediterranean high mountain ecosystems.

Aims: The aims of this study were (1) to explore the variation in plant traits in relation to environmental variability; (2) to analyse the functional strategies of species; and (3) to assess the habitat constraints for the species in the study area.

Methods: We sampled the floristic composition of 76 1 m?×?1 m plots on five summits over 2,100 m above sea level in the mountains of the Sistema Central, Spain. Soil properties and temperature and grazing disturbance parameters were recorded. Eight plant traits were assessed in 21 species. Environmental variability and the co-variation of functional traits were analysed by RDA and PCA, respectively. Plant traits and environmental variability were related using fourth-corner analysis.

Results: Traits related to resource acquisition, such as leaf size and N concentration, varied with soil temperature and estimated summer water availability. Leaf dry matter content was found to be related to estimated water availability and soil pH. Seed mass was a factor of snow cover duration and water availability, and clonality to the duration of the vegetative period and estimated water availability. Grazing disturbance was related to the mean plant height of the species.

Conclusions: The results suggest that low temperatures, rather than water shortage, may be the principal limiting factor for resource acquisition in plants. Nevertheless species establishment is limited by water shortage during summer in these Mediterranean high mountain communities.     

New national and regional bryophyte records, 37

Abstract

We analyzed the form of the UV-absorption spectra and calculated the bulk UV-absorption capacity of the methanolic extracts (BUVACME) in 15 bryophytes (five liverworts and 10 mosses) from unshaded aquatic habitats of Tierra del Fuego (Argentina). BUVACME was obtained as the area under the absorbance curve (AUC) in several ranges of the UV band to allow comparison with the literature data. Both the spectra form and the AUC values depended on the species considered. The spectra showed either no, one or two defined peaks. The peaks were probably due to phenolic derivatives, which could act as both screening compounds and antioxidants. The different AUCs calculated were highly and significantly correlated, and thus it may be unimportant which of these is used. The AUC values of most liverworts were higher than those of most mosses and in particular, the liverworts Noteroclada confluens and Triandrophyllum subtrifidum showed much higher BUVACMEs than those analyzed in any other bryophyte. Thus, the accumulation of UV-absorbing compounds might often increase protection against UV radiation in liverworts, but rarely in mosses. This is in line with the evolutionary differences between these two groups of bryophytes. Except for the two abovementioned liverwort species, BUVACME of aquatic bryophytes from Tierra del Fuego is not particularly different to that found in bryophytes from other zones of the planet.