Facilitation consequences for reproduction of the benefactor cushion plant Laretia acaulis along an elevational gradient: costs or benefits?

Environmental stress may favour facilitative interactions among plants but whether these interactions are positive for the benefactor and how this depends on stress factors, remains to be determined. We studied the effect of beneficiary cover and biomass on reproduction of the benefactor cushion plant Laretia acaulis (Apiaceae) in the central Chilean Andes during three years. Study sites were situated along an elevational gradient at 2600, 2800, 3000 and 3150 m a.s.l. This range comprises a cold‐ and a drought‐stress gradient, with moisture increasing and temperature decreasing with elevation. We studied the effect of natural gradients in beneficiary cover and of experimental cover removal on cushion flower and fruit production. Beneficiary cover had a negative effect on flower production but not on infructescence and fruit densities or fruit weights. A positive effect of beneficiaries on the fraction of flowers converted into fruits was detected for hermaphrodite cushions. The effect of beneficiary cover on flowering was independent of elevation or cushion gender, although these latter factors explained most of the variation. Removing the aboveground parts of the beneficiaries positively affected flowering at 2800 m a.s.l. but not at the other elevations. Our results suggest negative effects of facilitation on L. acaulis flowering, but these are neutralized in fruit production. Surprisingly, this conclusion holds along the entire elevational or stress gradient. This suggests that this system of facilitation is evolutionarily stable and not very sensitive to environmental change. It remains to be tested, however, whether facilitation affects fitness via growth and long‐term survival in these slow‐growing alpine cushions.     

Substantial compositional turnover of fungal communities in an alpine ridge‐to‐snowbed gradient

The main gradient in vascular plant, bryophyte and lichen species composition in alpine areas, structured by the topographic gradient from wind‐exposed ridges to snowbeds, has been extensively studied. Tolerance to environmental stress, resulting from wind abrasion and desiccation towards windswept ridges or reduced growing season due to prolonged snow cover towards snowbeds, is an important ecological mechanism in this gradient. The extent to which belowground fungal communities are structured by the same topographic gradient and the eventual mechanisms involved are less well known. In this study, we analysed variation in fungal diversity and community composition associated with roots of the ectomycorrhizal plant Bistorta vivipara along the ridge‐to‐snowbed gradient. We collected root samples from fifty B. vivipara plants in ten plots in an alpine area in central Norway. The fungal communities were analysed using 454 pyrosequencing analyses of tag‐encoded ITS1 amplicons. A distinct gradient in the fungal community composition was found that coincided with variation from ridge to snowbeds. This gradient was paralleled by change in soil content of carbon, nitrogen and phosphorus. A large proportion (66%) of the detected 801 nonsingleton operational taxonomic units (OTUs) were ascomycetes, while basidiomycetes dominated quantitatively (i.e. with respect to number of reads). Numerous fungal OTUs, many with taxonomic affinity to Sebacinales, Cortinarius and Meliniomyces, showed distinct affinities either to ridge or to snowbed plots, indicating habitat specialization. The compositional turnover of fungal communities along the gradient was not paralleled by a gradient in species richness.     

Shrub facilitation drives tree establishment in a semiarid fog‐dependent ecosystem

Questions

The exceptional occurrence of tall rain forest patches on foggy coastal mountaintops, surrounded by extensive xerophytic shrublands, suggests an important role of plant–plant interactions in the origin and persistence of these patches in semi‐arid Chile. We asked whether facilitation by shrubs can explain the growth and survival of rain forest tree species, and whether shrub effects depend on the identity of the shrub species itself, the drought tolerance of the tree species and the position of shrubs in regard to wind direction.

Location

Open area–shrubland–forest matrix, Fray Jorge Forest National Park, Chile.

Methods

We recorded survival after 12 years of a ~3600 tree saplings plantation (originally ~30‐cm tall individuals) of Aextoxicon punctatum, Myrceugenia correifolia and Drimys winteri placed outside forests, beneath the shrub Baccharis vernalis, and in open (shrub‐free) areas. We assessed the effects of neighbouring shrubs and soil humidity on survival and growth along a gradient related to the direction of fog movement.

Results

B. vernalis had a clear facilitative effect on tree establishment and survival since, after ~12 years, saplings only survived beneath the shrub canopy. Long‐term survival strongly depended on tree species identity, drought tolerance and position along the soil moisture gradient, with higher survival of A. punctatum (>35%) and M. correifolia (>14%) at sites on wind‐ and fog‐exposed shrubland areas. Sites occupied by the shrub Aristeguietia salvia were unsuitable for trees, presumably due to drier conditions than under B. vernalis.

Conclusions

Interactions between shrubs and fog‐dependent tree species in dry areas revealed a strong, long‐lasting facilitation effect on planted tree's survival and growth. Shrubs acted as benefactors, providing sites suitable for tree growth. Sapling mortality in the shrubland interior was caused by lower soil moisture, the consequence of lower fog loads in the air and thus insufficient facilitation. While B. vernalis was a key ecosystem engineer (nurse) and intercepted fog water that dripped to trees planted underneath, drier sites with A. salvia were unsuitable for trees. Consequently, nurse effects related to water input are strongly site and species specific, with facilitation by shrubs providing a plausible explanation for the initiation of forest patches in this semi‐arid landscape.     

Inhibitory effects of Eucalyptus globulus on understorey plant growth and species richness are greater in non‐native regions

Aim

We studied the novel weapons hypothesis in the context of the broadly distributed tree species Eucalyptus globulus. We evaluated the hypothesis that this Australian species would produce stronger inhibitory effects on species from its non‐native range than on species from its native range.

Location

We worked in four countries where this species is exotic (U.S.A., Chile, India, Portugal) and one country where it is native (Australia).

Time period

2009–2012.

Major taxa studied

Plants.

Methods

We compared species composition, richness and height of plant communities in 20 paired plots underneath E. globulus individuals and open areas in two sites within its native range and each non‐native region. We also compared effects of litter leachates of E. globulus on root growth of seedlings in species from Australia, Chile, the U.S.A. and India.

Results

In all sites and countries, the plant community under E. globulus canopies had lower species richness than did the plant community in open areas. However, the reduction was much greater in the non‐native ranges: species richness declined by an average of 51% in the eight non‐native sites versus 8% in the two native Australian sites. The root growth of 15 out of 21 species from the non‐native range were highly suppressed by E. globulus litter leachates, whereas the effect of litter leachate varied from facilitation to suppression for six species native to Australia. The mean reduction in root growth for Australian plants was significantly lower than for plants from the U.S.A., Chile and India.

Main conclusions

Our results show biogeographical differences in the impact of an exotic species on understorey plant communities. Consistent with the novel weapons hypothesis, our findings suggest that different adaptations of species from the native and non‐native ranges to biochemical compounds produced by an exotic species may play a role in these biogeographical differences.     

Trends in soil characteristics along a recently deglaciated foreland on Anvers Island,Antarctic Peninsula

We assessed patterns in soil development at a recently deglaciated foreland on Anvers Island on the Antarctic Peninsula. Soil samples were collected along transects extending 35 m over bare ground from the edge of a receding glacier; the far end of these transects has been ice free for approximately 20 years. We also compared soils at the far end of these transects under bare ground to those under canopies of isolated individuals of Deschampsia antarctica, a caespitose grass, that had recently colonized the site (established for <6 years). In addition, we compared soils at this young foreland to those in a well-developed tundra island that has been ice free for at least several hundred years. At the foreland site, soil moisture was greatest near the glacier, consistent with proximity to meltwater, and declined with distance from the glacier. This decline in soil moisture may explain the decrease in litter decomposition rates and the greater soil nitrate (NO₃ ⁻) concentrations that we observed with distance from the glacier. The greater soil moisture near the glacier likely promoted leaching and transport of NO₃ ⁻ to drier soils away from the glacier. The presence of D. antarctica at the glacier foreland had little effect on soil properties, which is not surprising considering it had only colonized sampling areas during the previous 5 years. Compared to the foreland, which contained only mineral soil, soil at the older tundra site had a 2.5- to 5-cm-thick organic horizon that had much higher concentrations of total carbon, nitrogen, and NO₃ ⁻.     

Correspondence of seed traits with niche position in glacier foreland succession

Changes in species composition during succession are driven by biotic and abiotic factors leading to a multitude of niches occupied by distinct species. Gradient analyses of plant communities provide opportunities to approximate the niche position of species along a successional gradient. Several plant traits have been used to explain mechanisms governing successional sequences, but generalising changes in species traits during primary succession is still controversial. This study examined whether the seed mass and the optimum temperature for germination could explain the niche position of several glacier foreland species along a primary successional gradient in the Austrian Central Alps. We hypothesised that pioneer species should possess lighter seeds and a lower optimum temperature for germination than late successional species. We found significant differences in the seed mass between species, but the seed mass did not correspond with the assigned niche position on the successional gradient. Germination responses to temperature also differed significantly between species. Pioneer species performed better at lower temperatures than late successional species, suggesting that the optimum temperature for germination is a driver of niche separation. We discuss the interactions between seed traits and environmental conditions along the primary successional gradient emphasising the importance of temperature requirements for the germination. Differences in the regeneration characteristics are a major cue governing species turnover in glacier foreland succession.     

Population dynamics along a primary succession gradient: do alpine species fit into demographic succession theory?

Background and Aims

Understanding processes and mechanisms governing changes in plant species along primary successions has been of major importance in ecology. However, to date hardly any studies have focused on the complete life cycle of species along a successional gradient, comparing pioneer, early and late-successional species. In this study it is hypothesized that pioneer species should initially have a population growth rate, λ, greater than one with high fecundity rates, and declining growth rates when they are replaced by late-successional species. Populations of late-successional species should also start, at the mid-successional stage (when pioneer species are declining), with growth rates greater than one and arrive at rates equal to one at the late successional stage, mainly due to higher survival rates that allow these species to persist for a long time.

Methods

The demography of pioneer- (Saxifraga aizoides), early (Artemisia genipi) and late-successional species (Anthyllis vulneraria ssp. alpicola) was investigated together with that of a ubiquitous species (Poa alpina) along the Rotmoos glacier foreland (2300–2400 m a.s.l., Central Alps, Austria) over 3 years. A matrix modelling approach was used to compare the main demographic parameters. Elasticity values were plotted in a demographic triangle using fecundity, individual growth and survival as vital rates contributing to the population growth rates.

Key Results

The results largely confirmed the predictions for population growth rates during succession. However, high survival rates of larger adults characterized all species, regardless of where they were growing along the succession. At the pioneer site, high mortality rates of seedlings, plantlets and young individuals were recorded. Fecundity was found to be of minor relevance everywhere, but it was nevertheless sufficient to increase or maintain the population sizes.

Conclusions

Demographically, all the species over all sites behaved like late-successional or climax species in secondary successions, mainly relying on survival of adult individuals. Survival serves as a buffer against temporal variation right from the beginning of the primary succession, indicating a major difference between primary and secondary succession.Key words: Demography, elasticity, glacier foreland, matrix model, population growth, primary succession, strategy, Saxifraga aizoides, Artemisia genipi, Anthyllis vulneraria ssp. alpicola, Poa alpina     

Can the biomass-ratio hypothesis predict mixed-species litter decomposition along a climatic gradient?

Background and Aims

The biomass-ratio hypothesis states that ecosystem properties are driven by the characteristics of dominant species in the community. In this study, the hypothesis was operationalized as community-weighted means (CWMs) of monoculture values and tested for predicting the decomposition of multispecies litter mixtures along an abiotic gradient in the field.

Methods

Decomposition rates (mg g⁻¹ d⁻¹) of litter from four herb species were measured using litter-bed experiments with the same soil at three sites in central France along a correlated climatic gradient of temperature and precipitation. All possible combinations from one to four species mixtures were tested over 28 weeks of incubation. Observed mixture decomposition rates were compared with those predicted by the biomass-ratio hypothesis. Variability of the prediction errors was compared with the species richness of the mixtures, across sites, and within sites over time.

Key Results

Both positive and negative prediction errors occurred. Despite this, the biomass-ratio hypothesis was true as an average claim for all sites (r = 0·91) and for each site separately, except for the climatically intermediate site, which showed mainly synergistic deviations. Variability decreased with increasing species richness and in less favourable climatic conditions for decomposition.

Conclusions

Community-weighted mean values provided good predictions of mixed-species litter decomposition, converging to the predicted values with increasing species richness and in climates less favourable to decomposition. Under a context of climate change, abiotic variability would be important to take into account when predicting ecosystem processes.     

Arctic root‐associated fungal community composition reflects environmental filtering

There is growing evidence that root‐associated fungi have important roles in Arctic ecosystems. Here, we assess the diversity of fungal communities associated with roots of the ectomycorrhizal perennial herb Bistorta vivipara on the Arctic archipelago of Svalbard and investigate whether spatial separation and bioclimatic variation are important structuring factors of fungal community composition. We sampled 160 plants of B. vivipara from 32 localities across Svalbard. DNA was extracted from entire root systems, and 454 pyrosequencing of ITS1 amplicons was used to profile the fungal communities. The fungal communities were predominantly composed of Basidiomycota (55% of reads) and Ascomycota (35%), with the orders Thelephorales (24%), Agaricales (13.8%), Pezizales (12.6%) and Sebacinales (11.3%) accounting for most of the reads. Plants from the same site or region had more similar fungal communities to one another than plants from other sites or regions, and sites clustered together along a weak latitudinal gradient. Furthermore, a decrease in per‐plant OTU richness with increasing latitude was observed. However, no statistically significant spatial autocorrelation between sites was detected, suggesting that environmental filtering, not dispersal limitation, causes the observed patterns. Our analyses suggest that while latitudinal patterns in community composition and richness might reflect bioclimatic influences at global spatial scales, at the smaller spatial scale of the Svalbard archipelago, these changes more likely reflect varied bedrock composition and associated edaphic factors. The need for further studies focusing on identifying those specific bioclimatic and edaphic factors structuring root‐associated fungal community composition at both global and local scales is emphasized.     

Patterns of Primary Succession on the Foreland of Coleman Glacier,Washington, USA

Patterns of community development vary among studied glacier forelands around the world. However, there have been few studies of primary succession on glacial forelands in temperate regions of North America. We described patterns in community composition, vegetation cover, diversity, and vegetation heterogeneity during primary succession on the foreland of Coleman Glacier, in Washington State, USA. Community composition changed rapidly with high turnover between age classes. Cover increased through succession as expected. Species richness and diversity were highest in early succession at small scales and in late succession at larger scales. At small scales, heterogeneity decreased in early succession but increased in mature sites. At larger scales, heterogeneity reached its lowest point earlier in succession. These scale-dependent patterns in diversity and heterogeneity differ from results of other studies of glacier forelands. We hypothesize that these patterns arise due to the development of a dense canopy of the deciduous shrub Alnus viridis followed by a dense canopy of Abies amabilis, Tsuga heterophylla, and Pseudotsuga menziesii.     

Fine nurse variations explain discrepancies in the stress‐interaction relationship in alpine regions

Despite a large consensus on increasing facilitation among plants with increasing stress in alpine regions, a number of different outcomes of interaction have been observed, which impedes the generalisation of the ‘stress‐gradient hypothesis’ (SGH). With the aim to reconcile the different viewpoints on the stress‐interaction relationship in alpine environments we hypothesized that fine nurse variations within a single life form (cushion) may explain this pattern variability. To test this hypothesis, we compared the magnitude of the stress‐interaction relationship in a single study area with that observed in existing studies involving cushions, worldwide. We characterized the nurse effects of cushions on the whole plant community at inter‐specific, intra‐specific and intra‐individual levels along a stress gradient in the dry, alpine tropics of Bolivia (4400 m, 4700 m and 4900 m a.s.l). Using a relative index of interaction (RII) we included our data in a meta‐analysis on the nurse effects of cushions along alpine gradients, worldwide. At inter‐specific level, the loose cushion Pycnophyllum was a better nurse than the compact Azorella compacta. However, at intra‐individual level facilitation was higher at the periphery than at the centre of cushions, exceeding in magnitude the variation observed at inter‐specific level. This pattern was associated with higher minimum temperature and lower mortality at the periphery of cushions. The net effects of cushions on plant communities became more positive at higher elevation, corroborating the SGH. Within our single site in Bolivia, fine morphological nurse variations captured a similar variability in the stress‐interaction relationship as that observed in a subset of studies on cushions on a worldwide scale. This suggests that fine variations in nurse traits, in general those not considered in protocols dealing with facilitation or in restoration/conservation management plans, explain in part the current discrepancies among SGH studies in alpine regions.     

The importance of facilitation in the zonation of shrubs along a coastal salinity gradient

Question: What are the interactive roles of abiotic stress and plant interactions in mediating the zonation of the shrub Tamarix chinensis along a salinity gradient? Location: Yellow River estuary (37°46′N, 119°09′E), northeast China. Methods: We surveyed the zonation of T. chinensis along a salinity gradient and quantified its salt tolerance using a pot experiment. In two field experiments, we transplanted T. chinensis seedlings into salt marsh, transitional zone and upland habitats, manipulated neighbours and quantified survivorship and biomass to examine neighbour effects. We also quantified vegetation effects on abiotic conditions in each zone. Results: Tamarix chinensis dominated the transitional zone, but was absent in upland and salt marsh habitats. In the pot experiment, T. chinensis grew well in freshwater treatments, but was inhibited by increasing salinity. Field experiments revealed that competition from neighbours limited T. chinensis growth in the uplands, while T. chinensis transplants were limited, with or without neighbours, in the salt marsh by high soil salinity. In the transitional zone, however, T. chinensis transplants performed better with than without neighbours. Vegetation removal significantly elevated soil salinity in the transitional zone, but not in other zones. Conclusions: Competition, facilitation and abiotic stress are all important in mediating the zonation of T. chinensis. Within its physiological stress tolerance range, or fundamental niche, it is limited by plant competition in low salinity habitats, and facilitated by neighbours in high salt stress habitats, but cannot survive in salt marshes having salinities above its salt stress tolerance limit. Our results have implications for understanding the relationships between facilitation and stress gradients.     

Changes in the structure and heterogeneity of vegetation and microsite environments with the chronosequence of primary succession on a glacier foreland in Ellesmere Island, high arctic Canada

Primary plant succession was investigated on a well-vegetated glacier foreland on Ellesmere Island in high arctic Canada. A field survey was carried out on four glacier moraines differing in time after deglaciation to assess vegetation development and microsite modification in the chronosequence of succession. The results showed evidence of directional succession without species replacement, which is atypical in the high arctic, reflecting the exceptionally long time vegetation development. During this successional process, Salix arctica dominated throughout all moraines. The population structures of S. arctica on these moraines implied the population growth of this species with progressing succession. The population density of S. arctica reflected the abundance of vascular plants, suggesting that development of the plant community might be related to structural changes and the growth of constituting populations. Through such growths of the population and the whole community with progressing succession, the spatial heterogeneity of vegetation gradually declines. Moreover, this vegetation homogenization is accompanied by changes in the spatial heterogeneity of microsite environments, suggesting significant plant effects on the modification of microsite environments. Accordingly, it was concluded that the directional primary succession observed on this glacier foreland is characterized by the initial sporadic colonization of plants, subsequent population growths, and the community assembly of vascular plants, accompanied by microsite modification.     

Local trampling disturbance effects on alpine plant populations and communities: Negative implications for climate change vulnerability

Global change is modifying species communities from local to landscape scales, with alterations in the abiotic and biotic determinants of geographic range limits causing species range shifts along both latitudinal and elevational gradients. An important but often overlooked component of global change is the effect of anthropogenic disturbance, and how it interacts with the effects of climate to affect both species and communities, as well as interspecies interactions, such as facilitation and competition. We examined the effects of frequent human trampling disturbances on alpine plant communities in Switzerland, focusing on the elevational range of the widely distributed cushion plant Silene acaulis and the interactions of this facilitator species with other plants. Examining size distributions and densities, we found that disturbance appears to favor individual Silene growth at middle elevations. However, it has negative effects at the population level, as evidenced by a reduction in population density and reproductive indices. Disturbance synergistically interacts with the effects of elevation to reduce species richness at low and high elevations, an effect not mitigated by Silene. In fact, we find predominantly competitive interactions, both by Silene on its hosted and neighboring species and by neighboring (but not hosted) species on Silene. Our results indicate that disturbance can be beneficial for Silene individual performance, potentially through changes in its neighboring species community. However, possible reduced recruitment in disturbed areas could eventually lead to population declines. While other studies have shown that light to moderate disturbances can maintain high species diversity, our results emphasize that heavier disturbance reduces species richness, diversity, as well as percent cover, and adversely affects cushion plants and that these effects are not substantially reduced by plant–plant interactions. Heavily disturbed alpine systems could therefore be at greater risk for upward encroachment of lower elevation species in a warming world.     

Carbohydrate reserves in the facilitator cushion plant <Emphasis Type="Italic">Laretia acaulis</Emphasis> suggest carbon limitation at high elevation and no negative effects of beneficiary plants

The elevational range of the alpine cushion plant Laretia acaulis (Apiaceae) comprises a cold upper extreme and a dry lower extreme. For this species, we predict reduced growth and increased non-structural carbohydrate (NSC) concentrations (i.e. carbon sink limitation) at both elevational extremes. In a facilitative interaction, these cushions harbor other plant species (beneficiaries). Such interactions appear to reduce reproduction in other cushion species, but not in L. acaulis. However, vegetative effects may be more important in this long-lived species and may be stronger under marginal conditions. We studied growth and NSC concentrations in leaves and stems of L. acaulis collected from cushions along its full elevational range in the Andes of Central Chile. NSC concentrations were lowest and cushions were smaller and much less abundant at the highest elevation. At the lowest elevation, NSC concentrations and cushion sizes were similar to those of intermediate elevations but cushions were somewhat less abundant. NSC concentrations and growth did not change with beneficiary cover at any elevation. Lower NSC concentrations at the upper extreme contradict the sink-limitation hypothesis and may indicate that a lack of warmth is not limiting growth at high-elevation. At the lower extreme, carbon gain and growth do not appear more limiting than at intermediate elevations. The lower population density at both extremes suggests that the regeneration niche exerts important limitations to this species’ distribution. The lack of an effect of beneficiaries on reproduction and vegetative performance suggests that the interaction between L. acaulis and its beneficiaries is probably commensalistic.     

Succession of terrestrial macrofungi along a deglaciation gradient at Glacier Blåisen, South Norway

A survey of terrestrial macrofungi along a deglaciation gradient from the glacier forefront to the established vegetation has been performed. The study was accomplished along a transect from Lake Finsevann to Glacier Blåisen, South Norway. Fortysix 6 m × 6 m plots divided into 9 subplots were positioned along this transect. The frequency of fruit bodies of each species in each plot was given on a 0–9 scale reflecting their occurrence in the subplots. Biotic and abiotic environmental variables for the plots were also recorded. The numerical methods used include principal component analysis (PCA), detrended correspondence analysis (DCA), and canonical correspondence analysis (CCA). Totally 99 species of macrofungi were recorded; 80.8 % of the species belonged to the Basidiomycota and 19.2 % to the Ascomycota. About one third of the species was only found once. In general, the species richness was highest on the glacier forefront, lowest in the most established vegetation. The proportion of species of Ascomycota was highest close to the glacier front. The main gradient in the data set was shown related to the primary ecological succession following the retreat of glaciers. The second gradient was possibly related to fine scale variation due to local moisture, microtopography, and secondary distribution. Of the variation in composition of species, 3 % can be ascribed to abiotic variance, 10 % to vascular plant variance, and 16 % to the combination of the abiotic and vascular plant component of variation. The results are discussed in the light of the time since deglaciation and the influence of certain biotic and abiotic environmental factors.     

Experimental test of the Invasional Meltdown Hypothesis: an exotic herbivore facilitates an exotic plant,but the plant does not reciprocally facilitate the herbivore

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