Overgrowth competition or facilitation from cushion plants: Implication for the role of plant-plant interactions.

Contrasting effects of two phenotypes of an alpine cushion plant on understory species drive community assembly

Cushion plants (as nurse species) can ameliorate soil conditions under their canopy in alpine environments. However, this amelioration (or the intensity of soil fertility islands) related with the size of plants is rarely studied. To assess size effects of Thylacospermum caespitosum on soil fertility islands, soil properties (pH; electric conductivity, EC; soil organic carbon, SOC; available nitrogen, AN; available phosphorus, AP; available potassium, AK) and microbial biomass (soil microbial biomass carbon, SMBC; soil microbial biomass nitrogen, SMBN) were investigated at three different elevations in the Qilian Mountains. Plant size significantly influenced (P < 0.001) soil properties (pH, EC, SOC, AN, AP, AK) and microbial biomass (SMBC and SMBN) at all three elevations. A size-dependent fertile island effect occurred beneath T. caespitosum, where the relative interaction index (RII) of soil properties and SMBC was affected (P < 0.001) by plant size at the three elevations. Moreover, most parameters of soil nutrition and microbial biomass under T. caespitosum were reduced (P < 0.001) with increases in elevation, but the RII was increased (P < 0.05). In short, soil amelioration by T. caespitosum was clearly dependent on plant size at all elevations, and this effect on soil increased with elevation. Thus, the existence of size-dependent fertility islands together with elevation should be regarded as a central mechanism of the nurse effect of T. caespitosum in harsh alpine ecosystems, where many ecological processes rely on the nurse effect of cushion plants.

One of the most important ecological processes is the formation of interspecific relationships in relation to spatial patterns among alpine cushion plants in extreme environmental habitats. However, such relationships remain poorly understood. Here, we examined the spatial patterns of alpine cushion plants along an altitudinal gradient of environmental severity and the interspecific relationship between two cushion species (Thylacospermum caespitosum and Androsace tangulashanensis) on the eastern Kunlun Mountain of China. Our results showed that the two species were highly aggregated within a distance of 2.5-5m at the mid (S2) altitude, whereas they were randomly distributed at the low (S1) and high (S3) altitudes. A positive spatial interaction between the two species was observed over shorter distances at the mid (S2) altitude, and the spatial patterns were related to the size of individuals of the two species. Moreover, the impact of A. tangulashanensis on T. caespitosum (RIIT. caespitosum) was negative in all the study plots, and a positive impact of T. caespitosum on A. tangulashanensis (RIIA. tangulashanensis) was only observed at the mid (S2) altitude. Together, these results demonstrated that the spatial patterns of these two cushions varied with environmental severity, since the outcome of the interactions were different, to some extent, at the three altitudes. Plant size is the main factor affecting the spatial correlation and interspecific relationship between two cushions. Therefore, its potential influence should be considered when discussing interspecific relationships among cushions and their community construction at small scales in alpine ecosystems.

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.

Functionally distinct assembly of vascular plants colonizing alpine cushions suggests their vulnerability to climate change.

As alpine plants, cushion species are particularly susceptible to environment changes. Thus, understanding population structure and community diversity variation of cushion plants along elevational gradients is crucial for estimating their response to predicted climate changes. In this study, Thylacospermum caespitosum populations from three elevations (low, medium, and high) in three climate zones of China (the Kunlun, Qilian, and Tianshan Mountains) were selected to evaluate the effect of elevation on the structure of T. caespitosum populations and species diversity of cushion communities. Results showed that elevation substantially influenced T. caespitosum populations (size structure, density, and death rate), as well as richness (α-diversity) and microhabitat species pool (species pool) of cushion communities. In the low elevations, T. caespitosum populations were in decline due to a lower ratio of small plants and higher mortality compared with populations at medium and high elevations. The α-diversity and species pool in cushion communities were significantly increased with decreased elevation, but the importance value of T. caespitosum decreased accordingly. Moreover, there was a significant positive correlation between elevation and relative importance value (the importance of one species in the community) of T. caespitosum (r = 0.883; P < 0.01). Elevation was significantly negatively correlated with the mortality rate of T. caespitosum (r = - 0.855; P < 0.01), α-diversity (r = - 0.933; P < 0.001), and species pool (r = - 0.885; P < 0.01). The declining characters of T. caespitosum population structure were obvious in low elevation populations. This decline may directly or indirectly relate to environmental change. Effects of elevation can provide an early indication of range contractions and population declines of cushion species with future climate warming. We call for more mechanistic studies of climate change impacts on cushion populations, particularly in alpine systems near the snow line.

Contrasting phenotypes of alpine cushion species have been recurrently described in several mountain ranges along small‐scale topography gradients, with tight competitive phenotypes in stressful convex topography and loose facilitative phenotypes in sheltered concave topography. The consistency of phenotypic effects along large‐scale climate stress gradients have been proposed as a test of the likely genetic bases of the differences observed at small‐scale. Inversely, plastic phenotypic effects are more likely to vanish at some points along climate stress gradients. We tested this hypothesis for two phenotypes of the alpine cushion species Thylacospermum caespitosum at four points along regional gradients of cold and drought stress in northwest China. We measured the traits of the two cushion phenotypes and quantified their associated plant communities and environmental variables along the regional temperature and aridity gradients. Cushion height, convexity and stem density overall showed significant effect of phenotypes. Difference in tightness of cushions between phenotypes was consistent across climate conditions, whereas differences in cushion convexity and height between phenotypes increased with increasing cold stress. Phenotypic effects on species richness and abundance were consistent along both climate gradients but not effects on species composition, while there were no phenotypic effects on environmental variables. Additionally, RII (relative interaction index) curves were linear along the drought gradient but unimodal along the temperature gradient, likely due to the occurrence of contrasting species pools at the different sites. We conclude that the consistency of phenotypic effects of T. caespitosum was high for species richness and abundance and mainly explained by differences in interference mediated by likely heritable differences in cushion tightness. Additionally, our study shows that the shapes of the relationship between plant responses to neighbours and environmental stresses are not necessarily driven by niche‐based deterministic factors.

垫状植物囊种草对群落物种多样性的影响

Summary In facilitative interactions, the beneficiary feedback effect (BFE) has been defined as the effect of beneficiary species (facilitated species) on their benefactor. BFEs have been shown to be dependent on environmental conditions and the composition of the beneficiary community. In alpine cushion systems, BFEs are more negative with more abundant, diverse and phylogenetically aggregated communities of beneficiary species. We tested the hypothesis that the functional composition of the beneficiary communities correlates with the direction and strength of BFE received by alpine cushion benefactors and specifically that a more negative BFE would occur with increasing density of graminoids and a more positive BFE would occur with increasing density of forbs and legumes. Additionally, we predicted that the negative BFE of graminoids would increase with increasing summer aridity. We used a data base of alpine cushion communities from 30 sites throughout the world to assess the overall relationship between the composition of beneficiary communities and the total flower density of cushion benefactors, and its variation with increasing drought. Additionally, in order to assess more precisely the role of the functional composition of the beneficiary communities on BFE in a very dry site with cushion benefactors exhibiting contrasting functional compositions of beneficiary communities, we also designed a field study in the Qilian Shan mountain range (China). At this site with a highly continental climate, we compared the number of flowers and fruits of different phenotypes of the alpine cushion species Thylacospermum caespitosum hosting numerous graminoids, numerous forbs or very few beneficiary species. In the intercontinental study, we found a negative relationship between graminoids and cushion benefactor flower density but no effect of other functional groups. The negative BFE of graminoids increased with increasing summer drought. In the dry Qilian Shan range, we found both a negative effect of graminoids on total flower density and a positive effect of forbs on flower density and fruit set. Our study indicates that the context dependence of BFE may be partially explained by the composition of beneficiary communities and in particular the negative effect of graminoids.

Field determination and QSPR prediction of equilibrium-status soil/vegetation partition coefficient of PCDD/Fs

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.

Cushions of Thylacospermum caespitosum (Caryophyllaceae) do not facilitate other plants under extreme altitude and dry conditions in the north-west Himalayas