Patterns of plant species turnover along grazing gradients

Questions: How are plant species distributed along grazing gradients? What is the shape of species richness patterns? How can we test for the existence of potential discontinuities in species turnover pattern? Location: Semi‐deserts in the eastern Caucasus, Azerbaijan, Gobustan district. Methods: We studied the distribution of vascular plant species along transects 900‐m long, perpendicular to five farms, and estimated grazing intensity as current livestock units per distance. We modelled species response curves with Huismann–Olff–Fresco (HOF) models and calculated species turnover by accumulating the first derivatives of all response curves. To test for potential discontinuities in changes of vegetation composition along the grazing gradient, we introduce a new null model based on the individualistic continuum concept that uses permutations of the observed pattern of species responses. Results: Most species show a sigmoidal negative response to grazing intensity, while a few species respond with a unimodal pattern. The monotonic decrease in species richness with increasing grazing intensity marks a process of overgrazing that leads to the complete extirpation of plant species. Although the species turnover pattern shows a clear peak, it does not deviate significantly from the null model of individualistic continuous changes. Conclusions: Our approach offers a method for differentiating between transition zones and continuous shifts in species composition along ecological gradients. It also provides a valuable tool for rangeland management to test state‐and‐transition concepts and gives deeper insights into ecological processes affected by grazing.     

Biodiversity effects on ecosystem functioning change along environmental stress gradients

Positive relationship between biodiversity and ecosystem functioning has been observed in many studies, but how this relationship is affected by environmental stress is largely unknown. To explore this influence, we measured the biomass of microalgae grown in microcosms along two stress gradients, heat and salinity, and compared our results with 13 published case studies that measured biodiversity–ecosystem functioning relationships under varying environmental conditions. We found that positive effects of biodiversity on ecosystem functioning decreased with increasing stress intensity in absolute terms. However, in relative terms, increasing stress had a stronger negative effect on low‐diversity communities. This shows that more diverse biotic communities are functionally less susceptible to environmental stress, emphasises the need to maintain high levels of biodiversity as an insurance against impacts of changing environmental conditions and sets the stage for exploring the mechanisms underlying biodiversity effects in stressed ecosystems.     

Plant size influences abundance of floral visitors and biomass allocation for the cushion plant Thylacospermum caespitosum under an extreme alpine environment

Variation in size may influence the abundance of visitors and reproductive allocation for cushion plants in the extreme alpine environments. To assess effects of plant size on the abundance of main visitors and reproductive allocation in Thylacospermum caespitosum populations at two altitudes, the abundance of the visitors, visiting frequency, total number of flowers, number of fruits, number of unseeded flowers, and reproductive allocation were investigated during the period of reproductive growth. Concurrently, the effects of plant size on the visitors' contributions to fruit setting rate were assessed by a bagging experiment. Our results showed that flies (Musca domestica and Dasyphora asiatica) were the main pollinating insects of T. caespitosum, and they could obvious facilitate (p < 0.05) the fruit setting rate of this cushion plant. Seed set and floral visitation were significantly influenced (p < 0.001) by plant size. Moreover, the reproductive allocation and fruit setting rate of T. caespitosum was influenced (p < 0.001) by plant size. More biomass was allocated to reproduction in plants of greater diameter. There is an increase in reproductive success (increases of fruit number with increase in plant size) in relation to plant size. In conclusion, the extent of M. domestica and D. asiatica to facilitate the fruit setting rate mainly depended on the size of T. caespitosum. Size‐dependent reproductive allocation occurred in T. caespitosum and was the chief factor affecting the contribution of flies to fruit setting rate. These traits reflect reproductive fitness of T. caespitosum related to plant size in extreme alpine environments.     

Variations in species and functional plant diversity along climatic and grazing gradients

Different components of biodiversity may vary independently of each other along environmental gradients giving insights into the mechanisms that regulate species coexistence. In particular, the functional diversity (FD) or the presence of rare or endemic species in natural assemblages do not necessarily increase with species diversity. We studied if different components of plant species diversity (species richness, Simpson diversity, evenness) varied similarly to FD (measured as a generalization of the Simpson index) and rarity along grazing intensity and climatic gradients. Plots under different sheep grazing regimes (high and low intensity, abandonment) were surveyed in five locations along a climatic gradient in north-eastern Spain, from semi-arid lowland to moist upland locations. Variation in species diversity, functional diversity and rarity followed different patterns. Species diversity was lowest in water-stressed environments (arid locations and southern aspects) and increased with grazing more makedly in humid locations. The FD was comparable between the most species-poor and species-rich locations and decreased with grazing in the moistest location, i.e. where species diversity markedly increased. The FD did not show a strong correlation with species richness nor with the Simpson index and less specious communities could show the highest functional diversity. The rarest species in the region were more frequently found in the abandoned areas, which held the lowest species diversity. Consequently, the mechanisms that enhance the diversity of species do not necessarily support a functional differentiation among those species or the maintenance of rare species in a region. We hypothesize that the degree of dependence of functional diversity on species diversity might be mostly related to the amplitude of the species' traits pool and on how species partition the niche space available.     

Projected impacts of climate change on regional capacities for global plant species richness

Climate change represents a major challenge to the maintenance of global biodiversity. To date, the direction and magnitude of net changes in the global distribution of plant diversity remain elusive. We use the empirical multi-variate relationships between contemporary water-energy dynamics and other non-climatic predictor variables to model the regional capacity for plant species richness (CSR) and its projected future changes. We find that across all analysed Intergovernmental Panel on Climate Change emission scenarios, relative changes in CSR increase with increased projected temperature rise. Between now and 2100, global average CSR is projected to remain similar to today (+0.3%) under the optimistic B1/+1.8°C scenario, but to decrease significantly (−9.4%) under the ‘business as usual’ A1FI/+4.0°C scenario. Across all modelled scenarios, the magnitude and direction of CSR change are geographically highly non-uniform. While in most temperate and arctic regions, a CSR increase is expected, the projections indicate a strong decline in most tropical and subtropical regions. Countries least responsible for past and present greenhouse gas emissions are likely to incur disproportionately large future losses in CSR, whereas industrialized countries have projected moderate increases. Independent of direction, we infer that all changes in regional CSR will probably induce on-site species turnover and thereby be a threat to native floras.     

Potential effects of climate change on plant species in the Faroe Islands

Aim To identify the effect of climate change on selected plant species representative of the main vegetation types in the Faroe Islands. Due to a possible weakening of the North Atlantic Current, it is difficult to predict whether the climate in the Faroe Islands will be warmer or colder as a result of global warming. Therefore, two scenarios are proposed. The first scenario assumes an increase in summer and winter temperature of 2 °C, and the second a decrease in summer and winter temperature of 2 °C. Location Temperate, low alpine and alpine areas in the northern and central part of the Faroe Islands. Methods The responses of 12 different plant species in the Faroe Islands were tested against measured soil temperature, expressed as T_min, T_max, snow cover and growing degree days (GDD), using generalised linear modelling (GLM). Results The tolerance to changes in winter soil temperature (0.3–0.8 °C) was found to be lower than the tolerance to changing summer soil temperature (0.7–1.0 °C), and in both cases lower than the predicted climate changes. Conclusions The species most affected by a warming scenario are those that are found with a limited distribution restricted to the uppermost parts of the mountains, especially Salix herbacea, Racomitrium fasciculare, and Bistorta vivipara. For other species, the effect will mainly be a general upward migration. The most vulnerable species are those with a low tolerance, especially Calluna vulgaris, and also Empetrum nigrum, and Nardus stricta. If the climate in the Faroe Islands should become colder, the most vulnerable species are those at low altitudes. A significantly lower temperature would be expected to produce a serious reduction in the extent of Vaccinium myrtillus and Galium saxatilis. Species like Empetrum nigrum, Nardus stricta, and Calluna vulgaris may also be vulnerable. In any case, these species can be expected to migrate downwards.     

Global patterns of genetic variation in plant species along vertical and horizontal gradients on mountains

Aim To understand global patterns of genetic variation in plant species on mountains and to consider the significance of mountains for the genetic structure and evolution of plant species. Location Global. Methods We review published studies. Results Genetic diversity within populations can vary along altitudinal gradients in one of four patterns. Eleven of 42 cited studies (26% of the total) found that populations at intermediate altitudes have greater diversity than populations at lower and higher altitudes. This is because the geographically central populations are under optimal environmental conditions, whereas the peripheral populations are in suboptimal situations. The second pattern, indicating that higher populations have less diversity than lower populations, was found in eight studies (19%). The third pattern, indicating that lower populations have lower diversity than higher populations, was found in 10 studies (24%). In 12 studies (29%), the intrapopulation genetic variation was found to be unaffected by altitude. Evidence of altitudinal differentiation was found in more than half of these studies, based on measurements of a range of variables including genome size, number of chromosomes or a range of loci using molecular markers. Furthermore, great variation has been found in phenotypes among populations at different altitudes in situ and in common garden experiments, even in cases where there was no associated variation in molecular composition. Mountains can be genetic barriers for species that are distributed at low elevations, but they can also provide pathways for species that occupy high‐elevation habitats. [Correction added after publication 9 October 2007: ‘less diversity’ changed to ‘greater diversity’ in the second sentence of the Results section of the Abstract] Main conclusions Genetic diversity within populations can vary along altitudinal gradients as a result of several factors. The results highlight the importance of phenotypic examinations in detecting altitudinal differences. The influence of mountain ridges on genetic differentiation varies depending, inter alia, on the elevation at which the species occurs. Based on these findings, zoning by altitudes or ridges would be helpful for the conservation of tree populations with the onset of global warming.     

Acceptability of plant species along grazing gradients around watering points in Tunisian arid zone

Water points provide excellent sites for studying overgrazing effects on plant communities in dry areas. Distance from water can be considered like a surrogate of grazing pressure being high near the water and low away from it. The main aim of this study is to investigate overgrazing effects on acceptability of fodder plants along a grazing gradient around three natural watering points. To achieve this goal, we classified spontaneous plants according to their acceptability degree and we followed their cover, richness and density as well as the grazing value along a grazing gradient around these wells, using phyto-ecological studies during the spring 2004 and 2006. Main results show that very palatable plants (mainly constituted by annuals) are more dominant in both the closed and the more disturbed transect areas around wells. The unpalatable plants dominate sites with moderate disturbance around wells. Ligneous palatable species obviously have a lower degree of disturbance. During the studied seasons the grazing gradient around wells 1 and 2, the oldest ones, seemed to exert a feedback upon the grazing intensity.     

Contrasting climate influences on Nothofagus pumilio establishment along elevational gradients

Predicted warmer temperatures and more frequent extreme climatic events in the southern Andes may affect the dynamics of the Patagonian forests. These environmental changes may differentially alter the probability of Nothofagus pumilio establishment across its altitudinal range of distribution. We monitored fruit fall, seedling emergence and survival at altitudinal distribution range of N. pumilio forests in Santa Cruz (49°22′ S—72°56′ W), Argentina. Fruit fall, seedling emergence and survival were tested in relation to drought, based on the SPEI (Standardized Precipitation Evapotranspiration Index), interacting with elevation. Fruit fall was consistently higher at low elevation and the abundance of fruits was not affected by drought intensity. Density of new-born seedlings was?~?9 to 24 times lower at low- than at high-elevation plots in our first observations (2014 and 2016), characterized by warm-dry climatic conditions in spring-early summers. In contrast, seedling abundance was?~?1.5 times larger at low elevations during relatively cold-wet growing seasons. Survival probability was explained by the interaction between SPEI and elevation. At low elevation and in dry periods, survival probability was lower (CI 54–72%) than in wet periods (CI 68–84%) but at high elevation similar survival was registered even with positive or negative SPEI values. Our results show interacting effects of elevation and drought on tree establishment at the elevation limits, with positive and negative drought effects at high and low elevations, respectively. Predicted increase of extreme drought events during the XXI century could be detrimental for N. pumilio establishment at dry, low-elevation forests.

     

Leaf damage by herbivore feeding guilds along gradients of elevation and plant species richness

The proportion of leaves damaged by specialist guilds such as miners and sap-feeders decreased at locations with higher species richness and at higher elevation in four subtropical forest seedling communities in south China. The effect of elevation was stronger in winter. Patterns of the generalist guild chewers were weaker.     

秦岭太白山木本植物物种多样性的梯度格局及环境解释

物种多样性沿环境梯度的分布格局是生物多样性研究的重要议题,而海拔梯度包含了各种环境因子的综合影响,因此对于探讨物种多样性沿环境梯度的分布格局具有重要意义。秦岭山脉地处我国暖温带向亚热带的过渡带,其植被垂直带和物种多样性分布格局对于研究我国南北部植被分异特征具有重要意义。基于对秦岭山脉太白山南北坡海拔1200—3750m之间的垂直样带调查的83个样方,本文利用植被数量分析方法(DCA和TWINSPAN)和Shannon-Wiener多样性指数、Pielou指数以及Jaccard相异性系数对太白山木本植物物种多样性在南北坡沿海拔梯度分布格局进行了初步研究。结果表明：太白山的木本植物群落具有明显的环境梯度格局,海拔是决定太白山植物群落分布的主要因素,而坡向起到次要作用一植物群落类型与坡向的关系不大,当考虑群落的环境梯度格局时,DCA第一轴主要与年均温密切相关,而第二轴则取决于年平均相对湿度：乔木层和灌木层的物种具有相似的海拔梯度格局,植物群落中木本植物物种丰富度和多样性随着海拔的升高单调下降;群落均匀度随着海拔变化的规律不明显;灌木层的物种多样性比乔木层更为丰富,而南坡具有比北坡更多的物种数和更高的多样性。相邻海拔之问群落的相异性在南北坡具有不同的分布格局,在北坡2800m以下,群落相异性沿海拔梯度变化不大,而在2800m以上的高海拔地区,群落相异性随海拔的升高而降低;在南坡,随着海拔的升高,群落相异性不断减少。太白山南坡群落比北坡分布更连续。     

Traits and climate are associated with first flowering day in herbaceous species along elevational gradients

Phenological responses to changing temperatures are known as “fingerprints of climate change,” yet these reactions are highly species specific. To assess whether different plant characteristics are related to these species‐specific responses in flowering phenology, we observed the first flowering day (FFD) of ten herbaceous species along two elevational gradients, representing temperature gradients. On the same populations, we measured traits being associated with (1) plant performance (specific leaf area), (2) leaf biochemistry (leaf C, N, P, K, and Mg content), and (3) water‐use efficiency (stomatal pore area index and stable carbon isotopes concentration). We found that as elevation increased, FFD was delayed for all species with a highly species‐specific rate. Populations at higher elevations needed less temperature accumulation to start flowering than populations of the same species at lower elevations. Surprisingly, traits explained a higher proportion of variance in the phenological data than elevation. Earlier flowering was associated with higher water‐use efficiency, higher leaf C, and lower leaf P content. In addition to that, the intensity of shifts in FFD was related to leaf N and K. These results propose that traits have a high potential in explaining phenological variations, which even surpassed the effect of temperature changes in our study. Therefore, they have a high potential to be included in future analyses studying the effects of climate change and will help to improve predictions of vegetation changes.     

Algal species diversity and dominance along gradients of stress and disturbance in marine environments

Data on algal species diversity from six areas along the Swedish coast, differing in salinity, length of growth period and grazing pressure were used to test two main predictions arising from the hump-backed model of species diversity (Grime 1973; Connell & Slatyer 1977; Tilman 1982; Fuentes & Jaksic 1988).Total number of species per m², total biomass per m² and primary production values for each species were calculated for 175 samples. Wave exposure was used as a measure of disturbance. The results are discussed in relation to stress (salinity and light) and disturbance factors (wave effects, ice scouring and grazing).A hump-backed model of species diversity, in relation to biomass per m² was found for all investigated areas. A similar response was also observed along local exposure gradients, with higher biomass and lower species numbers found at the sheltered sites compared to the more exposed ones. The most diverse communities were found at sites with intermediate wave exposure and in communities composed of species with intermediate primary production.It is suggested that the frequency of physical disturbance and severeness of stress strongly affects algal diversity and competition by determining the time interval over which successional replacement of species can occur. This lends support to the non-equilibrium view of community structure giving rise to humpbacked species diversity curves.     

Shifts in species richness, herbivore specialization, and plant resistance along elevation gradients

Environmental gradients have been postulated to generate patterns of diversity and diet specialization, in which more stable environments, such as tropical regions, should promote higher diversity and specialization. Using field sampling and phylogenetic analyses of butterfly fauna over an entire alpine region, we show that butterfly specialization (measured as the mean phylogenetic distance between utilized host plants) decreases at higher elevations, alongside a decreasing gradient of plant diversity. Consistent with current hypotheses on the relationship between biodiversity and the strength of species interactions, we experimentally show that a higher level of generalization at high elevations is associated with lower levels of plant resistance: across 16 pairs of plant species, low-elevation plants were more resistant vis-à-vis their congeneric alpine relatives. Thus, the links between diversity, herbivore diet specialization, and plant resistance along an elevation gradient suggest a causal relationship analogous to that hypothesized along latitudinal gradients.     

Testing the Stress-Gradient Hypothesis at the Roof of the World: Effects of the Cushion Plant Thylacospermum caespitosum on Species Assemblages

Many cushion plants ameliorate the harsh environment they inhabit in alpine ecosystems and act as nurse plants, with significantly more species growing within their canopy than outside. These facilitative interactions seem to increase with the abiotic stress, thus supporting the stress-gradient hypothesis. We tested this prediction by exploring the association pattern of vascular plants with the dominant cushion plant Thylacospermum caespitosum (Caryophyllaceae) in the arid Trans-Himalaya, where vascular plants occur at one of the highest worldwide elevational limits. We compared plant composition between 1112 pair-plots placed both inside cushions and in surrounding open areas, in communities from cold steppes to subnival zones along two elevational gradients (East Karakoram: 4850–5250 m and Little Tibet: 5350–5850 m). We used PERMANOVA to assess differences in species composition, Friedman-based permutation tests to determine individual species habitat preferences, species-area curves to assess whether interactions are size-dependent and competitive intensity and importance indices to evaluate plant-plant interactions. No indications for net facilitation were found along the elevation gradients. The open areas were not only richer in species, but not a single species preferred to grow exclusively inside cushions, while 39–60% of 56 species detected had a significant preference for the habitat outside cushions. Across the entire elevation range of T. caespitosum, the number and abundance of species were greater outside cushions, suggesting that competitive rather than facilitative interactions prevail. This was supported by lower soil nutrient contents inside cushions, indicating a resource preemption, and little thermal amelioration at the extreme end of the elevational gradient. We attribute the negative associations to competition for limited resources, a strong environmental filter in arid high-mountain environment selecting the stress-tolerant species that do not rely on help from other plants during their life cycle and to the fact the cushions do not provide a better microhabitat to grow in.