Enough space in a warmer world? Microhabitat diversity and small‐scale distribution of alpine plants on mountain summits

Aim

Global warming is predicted to shift distributions of mountain species upwards, driven by a release from climatic restrictions at their upper distribution limit and increased biotic pressure at their lower distribution limit. In alpine ecosystems, which are characterized by large microclimatic diversity and sparse vegetation cover, the relative importance of abiotic and biotic drivers for species distribution is poorly understood. To disentangle abiotic and biotic mechanisms affecting distributions of alpine species, we investigated how alpine plant species with differing elevational ranges and frequency trends over the past century differ in their microhabitat distribution, and how they respond to neighbouring vegetation.

Location

A total of 11 summits (2635—3410 m a.s.l.) in SE‐Switzerland.

Methods

We quantified the microscale abundance of 12 species in relation to biogeographic (frequency trend, i.e., change in occurrences over the past century, and elevational range on summits) and local microhabitat characteristics (temperature, substrate type). We assessed species size traits in relation to neighbouring vegetation characteristics to investigate possible neighbour interactions.

Results

Species with increasing frequency on summits over the past century were most abundant on scree and warmer slopes. Species with negative or stable frequency trends on summits were more abundant on organic soil and colder slopes. The preferred microhabitats of the latter were rarest overall, decreased with increasing elevation, and had the most competitive neighbours. Size of one high‐alpine specialist, Ranunculus glacialis was negatively related to cover of neighbouring vegetation, whereas other species showed no response to neighbours.

Main conclusions

Long‐term frequency trends of species correlate with their microhabitat association. Species with most negative frequency trends show preferences for the rarest microhabitat conditions, where they likely experience higher competitive pressure in a warming climate. This finding emphasizes the importance of characterizing microhabitat associations and microclimatic diversity to assess present and future distributions of alpine plant species.

     

Does plant diversity increase top–down control of herbivorous insects in tropical forest?

Higher trophic level interactions are key mediators of ecosystem functioning in tropical forests. A rich body of theory has been developed to predict the effects of plant diversity on communities at higher trophic levels and the mechanisms underlying such effects. The 'enemies hypothesis’ states that predators exert more effective top–down control of herbivorous insects with increasing plant diversity. Support for this hypothesis has been found in temperate forests and agroecosystems, but remains understudied in tropical forests. We compared incidence of attacks of different natural enemies using artificial caterpillars in a tropical forest landscape and investigated the role of plant community structure (i.e. species richness, composition and density), and the role of forest fragmentation (i.e. patch size, edge distance and canopy openness) on predation intensity. Plant community effects were tested with respect to three vegetation strata: trees, saplings and herbs. Observed predation was substantially due to ants. Predation rates increased with plant species richness for trees and herbs. Density of saplings, herb cover and herb species composition were important factors for predation. No significant patterns were found for fragmentation parameters, suggesting that forest fragmentation has not altered predation intensity. We conclude that in tropical forests, top–down control of herbivorous insects in the understory vegetation is affected by a combination of plant diversity, plant species composition and structural features of the plant community.     

Characteristics of nutrients in two dominant plant species and rhizospheric soils in alpine desert of the Qinghai-Xizang Plateau under contrasting climates北大核心CSCD

Aims This study was conducted to determine the responses of nutrients in plants and rhizospheric soils to climate in alpine-cold desert on the Qinghai-Xizang Plateau. Methods Tissue samples for two dominant plant species, Hippophae rhamnoides subsp. sinensis and Artemisia desertorum, and associated rhizospheric soil samples were collected from sites representing semi-Arid and sub-humid climates in the alpine-cold desert on the Qinghai-Xizang Plateau. Measurements were made on the contents of carbon, nitrogen and phosphorus in roots and shoots, as well as on organic carbon, total nitrogen, total phosphate, ammonium nitrogen, nitrate nitrogen and available phosphate in rhizospheric soils in the 0-10 cm and 10-20 cm layer. The relationship between nutrients in plant tissues and rhizospheric soils and the influencing factors were analyzed. Important findings There were significant differences between the semi-Arid and the sub-humid sites in tissue nutrients and rhizospheric soil nutrients for the two specie. Specifically, the contents of carbon, nitrogen, phosphorus in plant tissues differed significantly between the semi-Arid and the sub-humid sites. Soil organic carbon, total nitrogen, ammonium nitrogen, nitrate nitrogen and available phosphate for the rhizosphere of A. desertorum were significantly higher on site under sub-humid climate than that under semi-Arid climate; whereas the trend was reversed for the rhizosphere of H. rhamnoides subsp. sinensis. We found significant relationships between the tissue nutrients and soil nutrients, and significantly different plant nutrient ratios between the two species. There were negative correlations between tissues and rhizosheric soils in N:P ratio for A. desertorum and C:N ratio for H. rhamnoides subsp. sinensis under different climates. © 2018 Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

Do island plant populations really have lower genetic variation than mainland populations? Effects of selection and distribution range on genetic diversity estimates

Ecological and evolutionary studies largely assume that island populations display low levels of neutral genetic variation. However, this notion has only been formally tested in a few cases involving plant taxa, and the confounding effect of selection on genetic diversity (GD) estimates based on putatively neutral markers has typically been overlooked. Here, we generated nuclear microsatellite and plastid DNA sequence data in Periploca laevigata, a plant taxon with an island–mainland distribution area, to (i) investigate whether selection affects GD estimates of populations across contrasting habitats; and (ii) test the long‐standing idea that island populations have lower GD than their mainland counterparts. Plastid data showed that colonization of the Canary Islands promoted strong lineage divergence within P. laevigata, which was accompanied by selective sweeps at several nuclear microsatellite loci. Inclusion of loci affected by strong divergent selection produced a significant downward bias in the GD estimates of the mainland lineage, but such underestimates were substantial (>14%) only when more than one loci under selection were included in the computations. When loci affected by selection were removed, we did not find evidence that insular Periploca populations have less GD than their mainland counterparts. The analysis of data obtained from a comprehensive literature survey reinforced this result, as overall comparisons of GD estimates between island and mainland populations were not significant across plant taxa (N = 66), with the only exception of island endemics with narrow distributions. This study suggests that identification and removal of markers potentially affected by selection should be routinely implemented in estimates of GD, particularly if different lineages are compared. Furthermore, it provides compelling evidence that the expectation of low GD cannot be generalized to island plant populations.     

Historical plant records enlighten the conservation efforts of ferns and Lycophytes’ diversity in tropical China

Historical records, e.g., herbarium vouchers, contain information about species distribution since the early days of the scientific exploration of floras until today. These data provide crucial evidence to map the biodiversity of the area of interest and most importantly enable the evaluation of the conservation effectiveness for a given group of organisms. This study aimed to explore the ferns and lycophytes’ diversity of Xishuangbanna Dai Autonomous Prefecture in tropical China with special emphasis on conservation efforts provided by the currently established protected areas (PAs). Instead of relying exclusively on current observation, the database was compiled from digitalized herbarium vouchers and publications being explored with special attention on the temporal and spatial dimensions of collecting efforts. Utilizing the indices including species richness, weighed endemism, corrected weighted endemism, and beta diversity, hotspots of ferns and lycophytes’ diversity were identified. In turn, the proportion of hotspots located outside PAs was estimated as a measure of conservation gaps in Xishuangbanna. Our results revealed a long collecting history of ferns and lycophytes in Xishuangbanna and this prefecture accumulated a considerable number of historical records covering 20.2 % of Chinese and 3.6% of global fern diversity. The accumulation of historical records showed strong parallelism to the historical events shaping modern China. The spatial distribution of ferns and lycophytes in Xishuangbanna was characterized by a concentration of species richness in southern valleys and endemism in western and northern mountains. In terms of conservation, existing PAs showed higher effectiveness in the protection of species richness, whereas lower effectiveness was observed in the protection of endemism and beta diversity. Our research provided a key reference for understanding the diversity and conservation of ferns and lycophytes in Xishuangbanna, as well as highlighting the locality for future collecting and conservation efforts.     

Does nitrogen become limiting under high‐P conditions in detritus‐based tropical streams?

1. We examined effects of nutrients on leaf breakdown in interior forest streams at La Selva Biological Station, Costa Rica. We tested the hypothesis that dissolved inorganic nitrogen (DIN) becomes limiting when ambient phosphorus (P) concentration is high. We also compared the breakdown of relatively ‘low quality’ leaves (lower C : N, Trema integerrima) with that of ‘higher quality’ leaves (higher C : N, Ficus insipida) in a high‐P stream. 2. Litterbags were incubated in two streams: one enriched experimentally with P [target concentration 200 μg soluble reactive phosphorus (SRP) L^?1] and one control (naturally low P concentration approximately 10 μg SRP L^?1). Ammonium enrichment was achieved by adding fertiliser upstream of half of the litterbags in each stream. 3. Phosphorus addition stimulated leaf breakdown, microbial respiration, ergosterol and leaf %P. Leaf breakdown rate was consistent with those in La Selva streams with naturally high P concentration. 4. Nitrogen (N) addition had no effect on leaf breakdown, microbial respiration, ergosterol or leaf chemistry in either the P‐enriched or the reference stream, in spite of low N : P ratios. We conclude that N is probably not limiting in streams at La Selva that are naturally high in P. This may be due to moderately high ambient N concentration (>200 μg DIN L^?1) prevailing throughout the year. 5. The species with a lower C : N decomposed more rapidly and supported higher microbial activity than that with a higher C : N. Subtle differences in leaf N content, as well as dissolved P concentration, may be important in determining microbial colonisation and subsequent leaf breakdown.     

Does disturbance and restoration of alpine grassland soils affect the genetic structure and diversity of bacterial and N2-fixing populations?

Responses of bacterial communities to disturbance and restoration processes were investigated on alpine grassland soil. Bulk soil, rhizosphere soil and two soil separates, i.e. sand-size (2000-200 microm) and silt-size (50-2 microm) were sampled from undisturbed grassland soil to soil under restoration for 1 month, 1 year, 4 years and 13 years after disturbance. Automated ribosomal intergenic spacer analysis (ARISA) and restriction fragment length polymorphism (RFLP) of nifH gene pools were used to assay genetic structure of the bacterial communities and N2-fixing guild. According to the distribution of ARISA band length in bacterial phyla, the dominance of ARISA bands below 400 bp showed that Gram-positive bacteria would be predominant in the studied grassland soil when not disturbed. Disturbance affected the genetic structure of bacterial community and of N2-fixing guild in relation to their location within the selected habitats. Shifts in IGS and nifH profiles of bulk soil metagenome were larger than those observed from sand-size- and silt-size-fractions, accounting for 40-50% of the variance in the profiles. Restoration of the genetic structure of telluric bacteria community and N2-fixing populations was found to be influenced by the spatial heterogeneity of the soil and niche diversification. Particular bacterial genetic structure within distinct habitats were evidenced and must be defined as subdivisions of the meta-community of bulk soil. Scale of soil microbial diversity/stability relationships is discussed with special attention to disconnected bacterial habitat compared with whole soil with multiple niches.     

Does higher connectivity lead to higher genetic diversity? Effects of habitat fragmentation on genetic variation and population structure in a gypsophile

Habitat fragmentation is a major threat to the maintenance of genetic diversity in many plant populations. Genetic effects of population size have received far more attention than the effects of isolation—or connectivity—but both are key components of the fragmentation process. To analyze the consequences of fragment size and connectivity on the neutral genetic variation and population genetic structure of the dominant gypsophile Lepidium subulatum, we selected 20 fragments along two continuous gradients of size and degree of isolation in a fragmented gypsum landscape of Central Spain. We used eight polymorphic microsatellite markers, and analyzed a total of 344 individuals. Populations were characterized by high levels of genetic diversity and low inbreeding coefficients, which agrees with the mainly outcrossing system of L. subulatum and its high abundance in gypsum landscapes. Bayesian clustering methods, pairwise F _ST values and analysis of molecular variance revealed low among-population differentiation, with no significant isolation by distance. However, several genetic diversity indices such as allelic richness, number of effective alleles, expected heterozygosity and number of private alleles were negatively related to population isolation. The higher genetic diversity found on more connected fragments suggests higher rates of gene flow among more connected populations. Overall, our results highlight that fragmentation can have important effects on intra-population genetic processes even for locally abundant, dominant species. This, together with previously documented effects of connectivity on fitness of gypsophile species highlights the importance of including habitat connectivity in management and conservation strategies of this type of semiarid systems.     

Do pathogens reduce genetic diversity of their hosts? Variable effects of sylvatic plague in black‐tailed prairie dogs

Introduced diseases can cause dramatic declines in—and even the loss of—natural populations. Extirpations may be followed by low recolonization rates, leading to inbreeding and a loss of genetic variation, with consequences on population viability. Conversely, extirpations may create vacant habitat patches that individuals from multiple source populations can colonize, potentially leading to an influx of variation. We tested these alternative hypotheses by sampling 15 colonies in a prairie dog metapopulation during 7 years that encompassed an outbreak of sylvatic plague, providing the opportunity to monitor genetic diversity before, during and after the outbreak. Analysis of nine microsatellite loci revealed that within the metapopulation, there was no change in diversity. However, within extirpated colonies, patterns varied: In half of the colonies, allelic richness after recovery was less than the preplague conditions, and in the other half, richness was greater than the preplague conditions. Finally, analysis of variation within individuals revealed that prairie dogs present in recolonized colonies had higher heterozygosity than those present before plague. We confirmed plague survivorship in six founders; these individuals had significantly higher heterozygosity than expected by chance. Collectively, our results suggest that high immigration rates can maintain genetic variation at a regional scale despite simultaneous extirpations in spatially proximate populations. Thus, virulent diseases may increase genetic diversity of host populations by creating vacant habitats that allow an influx of genetic diversity. Furthermore, even highly virulent diseases may not eliminate individuals randomly; rather, they may selectively remove the most inbred individuals.     

When can refuges mediate the genetic effects of fire regimes? A simulation study of the effects of topography and weather on neutral and adaptive genetic diversity in fire‐prone landscapes

Understanding how landscape heterogeneity mediates the effects of fire on biodiversity is increasingly important under global changes in fire regimes. We used a simulation experiment to investigate how fire regimes interact with topography and weather to shape neutral and selection‐driven genetic diversity under alternative dispersal scenarios, and to explore the conditions under which microrefuges can maintain genetic diversity of populations exposed to recurrent fire. Spatial heterogeneity in simulated fire frequency occurred in topographically complex landscapes, with fire refuges and fire‐prone “hotspots” apparent. Interannual weather variability reduced the effect of topography on fire patterns, with refuges less apparent under high weather variability. Neutral genetic diversity was correlated with long‐term fire frequency under spatially heterogeneous fire regimes, being higher in fire refuges than fire‐prone areas, except under high dispersal or low fire severity (low mortality). This generated different spatial genetic structures in fire‐prone and fire‐refuge components of the landscape, despite similar dispersal. In contrast, genetic diversity was only associated with time since the most recent fire in flat landscapes without predictable refuges and hotspots. Genetic effects of selection driven by fire‐related conditions depended on selection pressure, migration distance and spatial heterogeneity in fire regimes. Allele frequencies at a locus conferring higher fitness under successional environmental conditions followed a pattern of “temporal adaptation” to contemporary conditions under strong selection pressure and high migration. However, selected allele frequencies were correlated with spatial variation in long‐term mean fire frequency (relating to environmental predictability) under weak dispersal, low selection pressure and strong spatial heterogeneity in fire regimes.     

Forecasting range shifts of a cold‐adapted species under climate change: are genomic and ecological diversity within species crucial for future resilience?

Cold‐adapted taxa are experiencing severe range shifts due to climate change and are expected to suffer a significant reduction of their climatically suitable habitats in the next few decades. However, it has been proposed that taxa with sufficient standing genetic and ecologic diversity will better withstand climate change. These taxa are typically more broadly distributed in geographic and ecological niche space, therefore they are likely to endure higher levels of populations loss than more restricted, less diverse taxa before the effects of those losses impact their overall diversity and resilience. Here, we explore the potential relationship between intraspecific genetic and ecological diversity and future resilience, using the cold‐adapted plant Primula farinosa. We employ high‐throughput sequencing to assess the genomic diversity of phylogeographic lineages in P. farinosa. Additionally, we use current climatic variables to define niche breadth and niche differentiation across lineages. Finally, we calibrate species distribution models (SDMs) and project the climatic preferences of each lineage on future climate to predict lineage‐specific shifts in climatically suitable habitats. Our study predicts relative persistence of future suitable habitats for the most genetically and ecologically diverse lineages of the cold‐adapted P. farinosa, but significant reduction of them for two out of its four lineages. While we do not provide specific experiments aimed at identifying the causal links between genetic diversity and resilience to climate change, our results indicate that greater genetic diversity and wider ecological breadth may buffer species responses to rapid climatic changes. This study further highlights the importance of integrating knowledge of intraspecific diversity for predicting species fate in response to climate change.     

Is plant temporal beta diversity of field margins related to changes in management practices?

Field margins have considerable ecological significance in agriculture-dominated landscapes by supporting biodiversity and associated services. However, agricultural changes during mid-20th century led to their drastic loss with a serious threat for biodiversity. Using time-series data, we aimed to get better insights into processes underlying plant patterns of field margins through time by i) quantifying plant temporal beta diversity components, ii) assessing whether the observed changes in plant communities can be related to changes in management practices applied to field margins. During the springs of 1994, 1998 and 2001, we surveyed plant communities and management practices of the same 116 field margins in three contrasted landscapes. We estimated temporal beta diversity in plant communities and partitioned it into its two dissimilarity resultant components, accounting for replacement of species (i.e. turnover) and for the nested gain or loss of species (i.e. nestedness). We then tested whether the observed changes in plant communities between 1994 and 1998 and, between 1998 and 2001 were related to changes in management practices using linear models. Plant communities of field margins exhibited strong temporal beta diversity dominated by turnover. Temporal turnover in plant communities was partly related to changes in management practices, i.e., a decrease of grazing concomitant to an increase of herbicide spraying. However, relationships were not consistent between all landscape contexts nor time period, suggesting that other unmeasured deterministic or stochastic processes could be driving the observed plant patterns. Taken together, our results suggest that maintaining a wide diversity of field margins with contrasted management contribute to maintaining plant diversity at a landscape scale. They underline the value of investigating plant temporal diversity patterns using time-series data and thus, the need to develop long-term studies making it possible to understand ecological processes shaping plant communities in agricultural landscapes.     

To what extent do microsatellite markers reflect genome‐wide genetic diversity in natural populations?

Microsatellite variability is widely used to infer levels of genetic diversity in natural populations. However, the ascertainment bias caused by typically selecting only the most polymorphic markers in the genome may lead to reduced sensitivity for judging genome-wide levels of genetic diversity. To test this potential limitation of microsatellite-based approaches, we assessed the degree of nucleotide diversity in noncoding regions of eight different carnivore populations, including inbred as well as outbred populations, by sequencing 10 introns (5.4–5.7 kb) in 20 individuals of each population (wolves, coyotes, wolverines and lynxes). Estimates of nucleotide diversity varied 30-fold (7.1 × 10⁻⁵ –2.1 × 10⁻³), with densities of one single nucleotide polymorphism every 112–5446 bp. Microsatellite genotyping (10–27 markers) of the same animals revealed mean multilocus heterozygosities of 0.54–0.78, a 1.4-fold difference among populations. There was a positive yet not perfect ( r ²  = 0.70) correlation between microsatellite marker heterozygosity and nucleotide diversity at the population level. For example, point estimates of nucleotide diversity varied in some cases with an order of magnitude despite very similar levels of microsatellite marker heterozygosity. Moreover, at the individual level, no significant correlation was found. Our results imply that variability at microsatellite marker sets typically used in population studies may not accurately reflect the underlying genomic diversity. This suggests that researchers should consider using resequencing-based approaches for assessing genetic diversity when accurate inference is critical, as in many conservation and management contexts.     

Can indirect selection and genetic context contribute to trait diversification? A transition‐probability study of blossom‐colour evolution in two genera

Darwin recognized that biological diversity has accumulated as a result of both adaptive and nonadaptive processes. Very few studies, however, have addressed explicitly the contribution of nonadaptive processes to evolutionary diversification, and no general procedures have been established for distinguishing between adaptive and nonadaptive processes as sources of trait diversity. I use the diversification of flower colour as a model system for attempting to identify adaptive and nonadaptive causes of trait diversification. It is widely accepted that variation in flower colour reflects direct, adaptive response to divergent selective pressures generated by different pollinators. However, diversification of flower colour may also result from the effects of nonadaptive, pleiotropic relationships with vegetative traits. Floral pigments that have pleiotropic relationships to vegetative pigments may evolve and diversify in at least two nonadaptive ways. (1) Indirect response to selection on the pleiotropically related nonfloral traits may occur (indirect selection). (2) Divergent evolution in response to parallel selective pressures (e.g. selection by pollinators for visually obvious flowers) may occur because populations are at different genetic starting points, and each population follows its own genetic `line of least resistance.' A survey of literature suggests that pleiotropic relationships between flower colour and vegetative traits are common. Phylogenetically informed analyses of comparative data from Dalechampia (Euphorbiaceae) and Acer (Aceraceae), based on trait‐transition probabilities and maximum likelihood, indicated that floral and vegetative pigments are probably pleiotropically related in these genera, and this relationship better explains the diversification of floral colour than does direct selection by pollinators. In Dalechampia pink/purple floral bract colour may have originated by indirect response to selection on stem and leaf pigments. In Acer selection by pollinators for visually obvious flowers may to have led to the evolution of red or purple flowers in lineages synthesizing and deploying red anthocyanins in leaves, and pale‐green or yellow flowers in species not deploying red anthocyanins in vegetative structures. This study illustrates the broader potential of indirect selection and parallel selection on different genetic starting points to contribute to biological diversity, and the value of testing directly for the operation of these nonadaptive diversifying processes.     

Are long‐unburnt eucalypt forest patches important for the conservation of plant species diversity?

Question: Are long‐unburnt patches of eucalypt forest important for maintaining floristic diversity? Location: Eucalyptus forests of southeastern New South Wales, Australia. Methods: Data from 976 sites representing a range of fire history from three major vegetation formations – shrubby dry sclerophyll forest (SF), grassy dry SF and wet SF – were analysed. Generalized linear models were used to examine changes in species richness with increasing time since wildfire and analysis of similarities to examine changes in community composition. Chi‐squared tests were conducted to examine the distribution of individual species across four time since fire categories. Results: Plant species relationships to fire varied between the three formations. Shrubby dry SF supported lower plant species richness with increasing time since wildfire and this was associated with shifts in community composition. Grassy dry SF showed significant shifts in community composition and species richness in relation to time, with a peak in plant species richness 20–30 yr post fire (either prescribed fire or wildfire). Wet SF increased in species richness until 10–20 yr post wildfire then displayed a general declining trend. Species richness in each vegetation type was not related to the fire frequencies and fire intervals observed in this study. Conclusions: Long‐unburnt (30–50 yr post wildfire) forests appeared to play a minor role in the maintenance of plant species diversity in dry forest systems, although this was more significant in wet forests. Maintenance of a range of fire ages within each vegetation formation will assist in maintaining floristic diversity within regions.     

What role do plant–soil interactions play in the habitat suitability and potential range expansion of the alpine dwarf shrub Salix herbacea?

Mountain plants may respond to warming climates by migrating along altitudinal gradients or, because climatic conditions on mountain slopes can be locally very heterogeneous, by migrating to different microhabitats at the same altitude. However, in new environments, plants may also encounter novel soil microbial communities, which might affect their establishment success. Thus, biotic interactions could be a key factor in plant responses to climate change. Here, we investigated the role of plant–soil feedback for the establishment success of the alpine dwarf shrub Salix herbacea L. across altitudes and late- and early snowmelt microhabitats. We collected S. herbacea seeds and soil from nine plots on three mountain-slope transects near Davos, Switzerland, and we transplanted seeds and seedlings to substrate inoculated with soil from the same plot or with soils from different microhabitats, altitudes and mountains under greenhouse conditions. We found that, on average, seeds from higher altitudes (2400–2700 m) and late-exposed snowbeds germinated better than seeds from lower altitudes (2200–2300 m) and early-exposed ridges. However, despite these differences in germination, growth was generally higher for plants from low altitudes, and there were no indications for a an home-soil advantage within the current range of S. herbacea. Interestingly, seedlings growing on soil from above the current altitudinal distribution of S. herbacea grew on average less well than on their own soil. Thus, although the lack of a home-soil advantage in the current habitat might be beneficial for S. herbacea in a changing environment, migration to habitats beyond the current altitudinal range might be limited, probably due to missing positive soil-feedback.