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1.
Abstract. Question: What are the tempo and mode of long‐term succession and of demographic processes in an alpine community, especially: tenacity, transition patterns, predictions, growth of individual cushion plants, cyclic succession, spatial patterns? Location: A low‐alpine mixed cushion /turf /snow‐tussock / shrub community in southeastern New Zealand. Methods: The distribution of seven plant cover‐types was recorded at 1024 fixed points in an 8 m × 8 m plot at approximately decade intervals for 50 years. The diameters of eight Donatia novae‐zelandiae cushions were monitored. Results: The process was essentially first‐order Markovian. There was a change in transition frequencies about 1980. The tenacity of the two major cover types — cushion and turf — was high, but that of cushion decreased about 1980 as some of its area was taken over by turf. The original informal prediction of 1955 that the cushion/turf would increase proved to be correct, probably because of paludification of the site. A prediction of 1987 made from observed transitions that cushion would dominate over turf has proved untrue because of a change in the transition probabilities in the 1980s, of unknown cause. There is a ten‐fold range in diameter growth rates among the eight cushions measured, but the mean rate of 5.3 mm.a‐1 is similar to that reported from other alpine and arctic sites. As cushions aged, turf colonized their centres, and in two cases new cushions colonized into this turf: consistent with cyclic succession. The pattern of transitions was compatible with a general interpretation of cyclic succession, but not definitive. Conclusions: Change is slow in this alpine community, and tenacity high. The change in transition frequencies about 1980, the invasion of individual cushions, and the decrease in spatial autocorrelation all suggest that cushions established on the site as a result of the clearance of woody vegetation after 1400 AD. Paludification may be causing some loss of tussock grass. A tendency for the cushions to break up, and the shallow peat accumulated below them, may indicate that they are the first generation of cushions on the site. Though these cushions are breaking up, other cushions are establishing, and cushions will continue to be an important part of the vegetation dynamics which may be part of a cyclic succession.  相似文献   

2.
Alpine ecosystems are among those biomes that are most vulnerable to climate change. Cushion plants are an important life form of alpine ecosystems and will likely play a critical role for the resilience of these habitats to climate change. We studied cushion size distribution and different measures of the compactness of cushions (biomass and rosette density, leaf area index) of the cushion plant, Androsace tapete along an elevational gradient from 4500 to 5200 m a.s.l. in the Nyainqentanglha Mountains of the central Tibetan Plateau. Cushion size distribution, total cover, and compactness of cushions varied substantially along the elevational gradient. At the driest site at low elevation we found the lowest total cushion cover, a particularly high proportion of very small cushions, and the most compact cushions (highest rosette and biomass densities, and leaf area index (LAI) per cushion). Our results indicate that in the semi‐arid Tibetan Plateau water availability is the more important climate factor than temperature affecting cushion plant traits and morphology.  相似文献   

3.
Arroyo  M.T.K.  Cavieres  L.A.  Peñaloza  A.  Arroyo-Kalin  M.A. 《Plant Ecology》2003,169(1):121-129
Low growing, compact cushion plants are a common and often dominant life form in temperate and subpolar alpine habitats. The cushion life-form can modify wind patterns, temperature and water availability and thus cushion species could be expected to act as nurse-plants facilitating the establishment of other alpine plant species on their surfaces. It has been suggested that the nurse effect should be most pronounced under more stressful environmental conditions, as found with increasing elevation in the alpine. One of the approaches used to detect the nurses has been the study of spatial associations among species, in which extreme clumping within or beneath one species has been interpreted as evidence of nursing. We characterized microclimatic conditions (soil and air temperature) within and outside cushions of Azorella monantha at two elevations (700 m a.s.l., corresponding to an elevation just above treeline, and 900 m a.s.l., corresponding to the upper limit of the cushion belt zone) on Cerro Diente in the Patagonian alpine of southern South America (50° S) and recorded all plant species growing upon cushions of various sizes and for paired sampling areas of equivalent sizes outside cushions. At 5 cm depth, soil temperature was slightly higher under cushions than under bare ground, but only significantly so at 900 m. Air temperature at ground level was significantly higher in the cushion microhabitat at both 700 m and 900 m, with the difference being more exaggerated at the highest elevation. At 700 m, a total of 27 species were recorded growing within cushions as compared to 29 outside cushions. At 900 m the corresponding numbers were 34 and 18. At the highest elevation, significantly more species grow within cushions than for equal areas outside cushions. Here moreover, 17 (48.6%) species grew preferentially within cushions, with eight of the latter being limited to the cushion microhabitat at this elevation. However, at 700 m there was no significant difference in species richness in the two microhabitats, and only one species (3.1%) grew preferentially on cushions. Considering individual species, nine occurring at both elevations showed non-preferential recruitment on cushions at 700 m, but significantly higher frequencies on cushions at 900 m. Results suggest striking altitudinal variation in the association with Azorella monantha on Cerro Diente, ranging from a very strong at 900 m to near absence at 700 m. Milder air and soil temperatures, shelter from wind, and greater water availability within cushions as opposed to outside cushions are discussed as possible factors favoring strong plant recruitment on cushions at higher elevations in the harsh Patagonian alpine environment. This revised version was published online in August 2006 with corrections to the Cover Date.  相似文献   

4.
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.  相似文献   

5.
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.  相似文献   

6.
Abstract. It has been proposed that in the harsh arctic and alpine climate zones, small microtopographic variations that can generate more benign conditions than in the surrounding environment could be perceived as safe sites for seedling recruitment. Cushion plants can modify wind pattern, temperature and water availability. Such modifications imply that cushion plants could act as ‘nurse plants’ facilitating the recruitment of other species in the community. This effect should be more evident under stressful conditions. We tested these hypotheses comparing the number of species that grow inside and outside Bolax gummifera cushions at two elevations (700 and 900 m a.s.l.) in the Patagonian Andes of Chile (50°S). At both elevations, and in equivalent areas, the number of species was registered within and outside cushions. A total of 36 and 27 plant species were recorded either within or outside B. gummifera cushions at 700 and 900 m a.s.l., respectively. At 700 m a.s.l., 33 species were recorded growing within cushions and 29 outside them, while at 900 m a.s.l. these numbers were 24 and 13 respectively. At both elevations there were significantly more species growing within than outside cushions, and the proportion of species growing within cushions increased with elevation. Thus there is a nurse effect of cushion plants and it is more evident at higher elevations. Shelter from wind and increased soil water availability seem to be the factors that increase plant recruitment within cushions.  相似文献   

7.
Question: Does the facilitative effect of cushion plants increase with elevation as a result of increases in environmental harshness? Does this hypothesis apply in the Sino‐Himalayan Mountains? Location: Lakaka Pass on the Baima Snow Mountains (28°20′N, 99°05′E), SW China. Methods: We evaluated the spatial association of several plant species with the cushion plant Arenaria polytrichoides (Caryophyllaceae) at two elevations (4500 m and 4700 m) in the study site and monitored temperature, moisture and nutritional status of soil beneath and outside the cushions. Results: While 14 species grow more frequently associated with the cushions at the higher elevation, at the lower site only three species were positively associated with cushions. Eleven of the species that occurred at both elevations changed their spatial association from neutral or negative with cushions at the lower site to positive at the higher elevation site. Substrate temperatures were rather similar between the cushions and areas of bare ground. Cushions maintained higher moisture than areas of bare ground at both elevations. Soils beneath cushions contained significantly more available nitrogen and potassium compared to open areas at the higher elevation. Conclusions: Our results show that facilitation by A. polytrichoides cushions increases with elevation in the Sino‐Himalayan region. This facilitation effect of A. polytrichoides cushions is probably due to the improved nutrient availability provided by cushion plants in the higher elevation, and these conditions probably permit increased plant recruitment, growth and survival.  相似文献   

8.
9.
Positive interactions between cushion plant and associated plants species in the high Andes of central Chile should also include the effects of fungal root symbionts. We hypothesized that higher colonization by arbuscular mycorrhizal (AM) fungi exists in cushion-associated (nursling) plants compared with conspecific individuals growing on bare ground. We assessed the AM status of Andean plants at two sites at different altitudes (3,200 and 3,600 m a.s.l.) in 23 species, particularly in cushions of Azorella madreporica and five associated plants; additionally, AM fungal spores were retrieved from soil outside and beneath cushions. 18 of the 23 examined plant species presented diagnostic structures of arbuscular mycorrhiza; most of them were also colonized by dark-septate endophytes. Mycorrhization of A. madreporica cushions showed differences between both sites (68% and 32%, respectively). In the native species Hordeum comosum, Nastanthus agglomeratus, and Phacelia secunda associated to A. madreporica, mycorrhization was six times higher than in the same species growing dispersed on bare ground at 3,600 m a.s.l., but mycorrhiza development was less cushion dependent in the alien plants Cerastium arvense and Taraxacum officinale at both sites. The ratio of AM fungal spores beneath versus outside cushions was also 6:1. The common and abundant presence of AM in cushion communities at high altitudes emphasizes the importance of the fungal root symbionts in such situations where plant species benefit from the microclimatic conditions generated by the cushion and also from well-developed mycorrhizal networks.  相似文献   

10.
11.
Question: In stressful abiotic environments positive plant interaction is expected to be a frequent and an important process driving community composition and structure. In the high Andes in central Chile, the cushion plant Azorella madreporica dominates plant communities and appears to benefit the assemblage of species that grows within it. However, there are also many other species that grow outside this nurse cushion plant, which may or may not interact with this species. What is the prevailing type of spatial associations among the plant species that are not growing inside the nurse plant? What is the type of interactions between cushion plants and those species growing outside them? Location: Molina River basin (33°20'S, 70°16’ W, 3600 m a.s.l.), in the Andes of central Chile, ca. 50 km east of Santiago. Methods: Two accurate mapping plots of individual plants of different species were located at two summits (Franciscano and Tres Puntas sites). The spatial distributions and associations between species growing outside cushions and within cushions at each site were estimated by point‐pattern analyses using the univariate and bivariate transformations of Ripley's K‐functions. Results: We found both positive and, especially, negative spatial associations (8 out of 12 species in Franciscano site) between A. madreporica cushions and plants growing outside them. However, most of the species showed positive spatial associations among them. The variation in spatial association was site‐specific and also depended on the type of plants involved. Adesmia spp., the second most abundant non‐cushion species, displayed negative associations with cushions and positive associations with other species growing outside cushions. Conclusions: Our study suggests very complex interactions among species, which ranged from positive to negative, and are also affected by abiotic environmental conditions.  相似文献   

12.

Background

The stress‐gradient hypothesis predicts a shift from facilitative to competitive plant interactions with decreasing abiotic stress. This has been supported by studies along elevation and temperature gradients, but also challenged by the hypothesis of a facilitation collapse at extremely harsh sites. Although facilitation is known to be important in primary succession, few studies have examined these hypotheses along primary succession gradients.

Aim

To examine whether there is a relationship between the presence of the circumpolar cushion plant Silene acaulis and other species, and if so, whether there is a shift between positive and negative interactions along a primary succession gradient in a glacier foreland.

Location

Finse, southern Norway.

Methods

We examined the performance of the common alpine forb Bistorta vivipara, species richness of vascular plants, bryophytes and lichens, and the number of seedlings and fertile vascular plants in S. acaulis cushions, and control plots without S. acaulis, along a succession gradient with increasing distance from a glacier front, and thus decreasing abiotic stress. To examine if S. acaulis cushions modify the abiotic environment, we recorded soil temperature, moisture, organic content and pH in cushions and control plots.

Results

Bistorta vivipara performed better, as shown by bigger leaves in S. acaulis cushions compared to control plots in the harshest part of the gradient close to the glacier. There were few differences in B. vivipara performance between cushion and control plots in the more benign environment further away from the glacier. This suggests a shift from facilitative to mainly neutral interactions by S. acaulis on the performance of B. vivipara with decreasing abiotic stress. A trend, although not significant, of higher vascular species richness and fertility inside S. acaulis cushions along the whole gradient, suggests that S. acaulis also facilitates community‐level species richness. The causal mechanism of this facilitation is likely that the cushions buffer extreme temperatures.

Conclusions

Our results support the stress‐gradient hypothesis for the relationship between the cushion plant S. acaulis and the performance of a single species along a primary succession gradient in a glacier foreland. S. acaulis also tended to increase vascular plant species richness and fertility regardless of stress level along the gradient, suggesting facilitation at the community level. We found no collapse of facilitation at the most stressful end of the gradient in this alpine glacier foreland.  相似文献   

13.
Seed weight is a prominent life history trait of plants affecting dispersal, establishment, and survival. In alpine environments, the few studies investigating the effect of elevation on seed weight within species have mainly detected a decrease in seed weight with increasing elevation. This relationship is generally attributed to the adverse climate at high elevations. In order to test this hypothesis, we analyzed seed weight variation across altitudes (2,435–3,055 m a.s.l.) in two consecutive years that differed in weather conditions in the high-alpine cushion plant Eritrichium nanum. We found a significant reduction in seed weight with increasing elevation in both years, but in the growing season with more adverse weather conditions, the reduction was more substantial than in the more favorable year. We conclude that alpine plants may be able to produce well-developed seeds at low elevations in almost all years, independent of weather conditions, whereas reproduction through seeds is potentially limited to years of favorable weather at high elevation.  相似文献   

14.
Ecosystem engineers are organisms able to modulate environmental forces and, hence, may change the habitat conditions for other species. In so doing, ecosystem engineers may affect both species richness and evenness of communities and, in consequence, change species diversity. If these changes in community attributes are related to the magnitude of the habitat changes induced by the engineers, it seems likely that engineer species will have greater effects on diversity in sites where they cause larger habitat changes. We addressed this issue by evaluating the effects of three alpine cushion plants on species richness, evenness, and diversity of high-Andean plant communities. Given that the difference in microclimatic conditions between cushions and the external environment increases with elevation, we proposed that these organisms should have greater effects on community attributes at higher than at lower elevation sites. Results showed that the three cushion species had positive effects on species richness, diversity, and evenness of plant communities. It was also observed that the magnitude of these effects changed with elevation: positive effects on species richness and diversity increased towards upper sites for the three cushions species, whereas positive effects on evenness increased with elevation for one cushion species but decreased with elevation for other two cushion species. These results suggest that the presence of cushions is important to maintain plant diversity in high-Andean communities, but this positive effect on diversity seems to increase as the difference in environmental conditions between cushions and the external environment increases with elevation.  相似文献   

15.
Abstract We developed multiple a priori hypotheses to link the observed spatial patterns with colonisation processes in the high alpine cushion plant, Azorella madreporica. We conducted this study in the Molina River basin (33°20′ S, 70°16′ W, 3600 m a.s.l.), in the Andes of central Chile, approximately 50 km east of Santiago. We mapped and measured size (as a surrogate for age) of individual cushions in two populations and used a standard spatial analytical tool (semivariograms) to test our alternative a priori hypotheses related to colonisation mode of the cushion species. In both populations, the size distribution of A. madreporica reflected a negative exponential or inverse‐J pattern, typical of uneven‐aged populations, where most of the cushions belonged to relatively smaller size classes, in effect, a regular success in the establishment of seedlings, where all size classes of cushions were represented in the population. The results were site‐specific, where best‐fit semivariograms for spatial cushion's size distribution suggested a gradual colonisation in one population and an episodic colonisation in the other population. Microsite distribution proved to be homogeneous at both sites. Thus, the study of the spatial explicit size‐age population distribution of an alpine species provides valuable information about the frequency, magnitude and site variation of the reproductive pulses in these harsh environments.  相似文献   

16.
Abstract: The habit of cushion growth positively affects plant temperature but at the same may increase the risk of occasional overheating. In order to determine the adaptive response to short-term heat stress, we exposed S. acaulis cushions at field sites to controlled heat treatments using infrared lamps. Natural diurnal changes in heat tolerance were monitored at alpine sites and at a site distinctly below the natural distribution boundary, where higher temperatures were expected. The range of heat tolerance limits in summer, 45.5 - 54.5 °C (9 K), exceeded that reported for other alpine species (0.1 - 5 K) and even that for total seasonal changes (5 - 8 K). Heat tolerance either increased or decreased on most days (80 %). The maximum diurnal increase was + 4.7 K. Under the experimental conditions heat hardening started at leaf temperatures around 30 °C and proceeded at mean rates of 1.0 ± 0.5 K/h. The onset of functional disturbances in photosystem II also occurred at 30 °C. Heating rates exceeding those naturally found above 30 °C (> 10 K/h) appeared to retard heat hardening. During summer average leaf temperature maxima were 12.4 K (600 m) and 13.0 K (1945 m) higher than air temperature which corroborates the heat trapping nature of cushion plants. At 600 m, as compared to 1945 m, cushions experienced significantly higher leaf temperature maxima (+ 8.8 K) and exceeded 30 °C on most days (80 %). This resulted in a significantly higher heat tolerance (LT50) at 600 m (51.7 ± 0.2 °C) than at 1945 m (49.8 ± 0.2 °C). The fast short-term changes of heat tolerance in summer help S. acaulis to cope with the occasional diurnal short-term heat stress associated with cushion growth.  相似文献   

17.
Climate change is driving species' range shifts, which are in turn disrupting species interactions due to species‐specific differences in their abilities to migrate in response to climate. We evaluated the consequences of asynchronous range shifts in an alpine plant–pollinator community by transplanting replicated alpine meadow turfs downslope along an elevational gradient thereby introducing them to warmer climates and novel plant and pollinator communities. We asked how these novel plant–pollinator interactions affect plant reproduction. We found that pollinator communities differed substantially across the elevation/temperature gradient, suggesting that these plants will likely interact with different pollinator communities with warming climate. Contrary to the expectation that floral visitation would increase monotonically with warmer temperatures at lower elevations, visitation rate to the transplanted communities peaked under intermediate warming at midelevation sites. In contrast, visitation rate generally increased with temperature for the local, lower elevation plant communities surrounding the experimental alpine turfs. For two of three focal plant species in the transplanted high‐elevation community, reproduction declined at warmer sites. For these species, reproduction appears to be dependent on pollinator identity such that reduced reproduction may be attributable to decreased visitation from key pollinator species, such as bumble bees, at warmer sites. Reproduction in the third focal species appears to be primarily driven by overall pollinator visitation rate, regardless of pollinator identity. Taken together, the results suggest climate warming can indirectly affect plant reproduction via changes in plant–pollinator interactions. More broadly, the experiment provides a case study for predicting the outcome of novel species interactions formed under changing climates.  相似文献   

18.
Determining which drivers lead to a specific species assemblage is a central issue in community ecology. Although many processes are involved, plant–plant interactions are among the most important. The phylogenetic limiting similarity hypothesis states that closely related species tend to compete stronger than distantly related species, although evidence is inconclusive. We used ecological and phylogenetic data on alpine plant communities along an environmental severity gradient to assess the importance of phylogenetic relatedness in affecting the interaction between cushion plants and the whole community, and how these interactions may affect community assemblage and diversity. We first measured species richness and individual biomass of species growing within and outside the nurse cushion species, Arenaria tetraquetra. We then assembled the phylogenetic tree of species present in both communities and calculated the phylogenetic distance between the cushion species and its beneficiary species, as well as the phylogenetic community structure. We also estimated changes in species richness at the local level due to the presence of cushions. The effects of cushions on closely related species changed from negative to positive as environmental conditions became more severe, while the interaction with distantly related species did not change along the environmental gradient. Overall, we found an environmental context‐dependence in patterns of phylogenetic similarity, as the interaction outcome between nurses and their close and distantly‐related species showed an opposite pattern with environmental severity.  相似文献   

19.
In alpine habitats, positive interactions among plants tend to increase with elevation as a result of altitudinal increase in environmental harshness. However, in mountains located in arid zones, lower elevations are also stressful because of scarce availability of water, suggesting that positive interactions may not necessarily increase with elevation. Here we analysed the spatial association of plant species with the nurse cushion plant Laretia acaulis at two contrasting elevations, and monitored the survival of seedlings of two species experimentally planted within and outside cushions in the semiarid Andes of central Chile. Positive spatial associations with cushions were more frequent at lower elevations. Species growing at the two elevations changed the nature of their association with cushions from neutral or negative at higher elevations to positive at lower elevations. Survival of seedlings was higher within cushions, particularly at lower elevations. The increased facilitation by cushions at lower elevations seems to be related to provision of moisture. This result suggests that cushion plants play a critical role in structuring alpine plant communities at lower elevations, and that climatic changes in rainfall could be very relevant for persistence of plant communities.  相似文献   

20.
Positive interactions among native plant species are common in alpine habitats, particularly those where one species (nurse plant) generates microclimatic conditions that are more benign than the surrounding environment, facilitating the establishment of other species. Nonetheless, these microclimatic conditions could facilitate the establishment of non-native species as well. A conspicuous component of the alien alpine flora of the central Chilean Andes is the perennial herb Taraxacum officinale agg. (dandelion). In contrast to other alien species that are restricted to human-disturbed sites, T. officinale is frequently observed growing within native plant communities dominated by cushion plants. In this study we evaluated if T. officinale is positively associated with the cushion plant Azorella monantha. Via seedling survival experiments and gas-exchange measurements we also assessed the patterns of facilitation between cushions and dandelions, and explore the potential mechanisms of invasion by dandelions. T. officinale grows spatially positively associated with cushions of A. monantha. Survival of seedlings, as well as their net-photosynthetic rates and stomatal conductance, were higher within cushions than in open areas away from them, suggesting that the microclimatic modifications generated by this native cushion facilitates the establishment and performance of a non-native invasive species. Our results, as well as other recent studies, highlight the role of native communities in facilitating rather than constraining non-native plant invasions, particularly in stressful habitats such as alpine environments.  相似文献   

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