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1.
《Plant Ecology & Diversity》2013,6(5-6):453-468
Background: In tundra ecosystems, the adjustment of phenological events, such as bud burst, to snowmelt timing is crucial to the climatic adaptation of plants. Natural small-scale variations in microclimate potentially enable plant populations to persist in a changing climate.

Aims: To assess how plant phenology responds to natural differences in snowmelt timing.

Methods: We observed the timing of eight vegetative and reproductive phenophases in seven dwarf-shrub species in relation to differences in snowmelt timing on a small spatial scale in an alpine environment in subarctic Finland.

Results: Some species and phenophases showed accelerated development with later snowmelt, thus providing full or partial compensation for the shorter snow-free period. Full compensation resulted in synchronous occurrence of phenophases across the snowmelt gradient. In other species, there was no acceleration of development. The timing of phenophases varied between two consecutive years and two opposing mountain slope aspects.

Conclusions: The results have shown three distinct patterns in the timing of phenophases in relation to snowmelt and suggest alternative strategies for adaptation to snowmelt timing. These strategies potentially apply to other species and tundra ecosystems and provide a framework, enabling one to compare and generalise phenological responses to snowmelt timing under different future climate scenarios.  相似文献   

2.
In alpine habitats, predicted warmer and longer growing seasons will influence plant phenology, with important implications for species adaptation and vegetation dynamics. However, little is known on the temperature sensitivity of different phenophases and on the characteristics allowing phenological variation among and within species. By integrating interannual micro‐climatic variability with experimental warming, we explored how the phenology of three alpine species is influenced by temperature and what mechanisms underlie intra‐ and inter‐specific phenological differences. The present study demonstrated that alpine plants have different temperature responses during their reproductive cycle, do not have constant thermal thresholds and heat‐use efficiencies to achieve the seed dispersal stage and can change their temperature sensitivity to flower along snowmelt gradients. In addition, the length of the reproductive cycle, which proved to be species‐specific under experimental warming, does not seem to be the only life‐history trait under selective pressure due to the short‐length of the snow‐free period. In a warming climate scenario, the phenology of sexual reproduction will be considerably altered, and alpine plants may be subjected to changes in population dynamics driven by altered perception of environmental cues appropriate for coordinating the timing of key life‐history events.  相似文献   

3.
Global climate change is already having significant impacts on arctic and alpine ecosystems, and ongoing increases in temperature and altered precipitation patterns will affect the strong seasonal patterns that characterize these temperature‐limited systems. The length of the potential growing season in these tundra environments is increasing due to warmer temperatures and earlier spring snow melt. Here, we compare current and projected climate and ecological data from 20 Northern Hemisphere sites to identify how seasonal changes in the physical environment due to climate change will alter the seasonality of arctic and alpine ecosystems. We find that although arctic and alpine ecosystems appear similar under historical climate conditions, climate change will lead to divergent responses, particularly in the spring and fall shoulder seasons. As seasonality changes in the Arctic, plants will advance the timing of spring phenological events, which could increase plant nutrient uptake, production, and ecosystem carbon (C) gain. In alpine regions, photoperiod will constrain spring plant phenology, limiting the extent to which the growing season can lengthen, especially if decreased water availability from earlier snow melt and warmer summer temperatures lead to earlier senescence. The result could be a shorter growing season with decreased production and increased nutrient loss. These contrasting alpine and arctic ecosystem responses will have cascading effects on ecosystems, affecting community structure, biotic interactions, and biogeochemistry.  相似文献   

4.
Climate change is altering spring snowmelt patterns in alpine and arctic ecosystems, and these changes may alter plant phenology, growth and reproduction. To predict how alpine plants respond to shifts in snowmelt timing, we need to understand trait plasticity, its effects on growth and reproduction, and the degree to which plants experience a home-site advantage. We tested how the common, long-lived dwarf shrub Salix herbacea responded to changing spring snowmelt time by reciprocally transplanting turfs of S. herbacea between early-exposure ridge and late-exposure snowbed microhabitats. After the transplant, we monitored phenological, morphological and fitness traits, as well as leaf damage, during two growing seasons. Salix herbacea leafed out earlier, but had a longer development time and produced smaller leaves on ridges relative to snowbeds. Longer phenological development times and smaller leaves were associated with reduced sexual reproduction on ridges. On snowbeds, larger leaves and intermediate development times were associated with increased clonal reproduction. Clonal and sexual reproduction showed no response to altered snowmelt time. We found no home-site advantage in terms of sexual and clonal reproduction. Leaf damage probability depended on snowmelt and thus exposure period, but had no short-term effect on fitness traits. We conclude that the studied populations of S. herbacea can respond to shifts in snowmelt by plastic changes in phenology and leaf size, while maintaining levels of clonal and sexual reproduction. The lack of a home-site advantage suggests that S. herbacea may not be adapted to different microhabitats. The studied populations are thus unlikely to react to climate change by rapid adaptation, but their responses will also not be constrained by small-scale local adaptation. In the short term, snowbed plants may persist due to high stem densities. However, in the long term, reduction in leaf size and flowering, a longer phenological development time and increased exposure to damage may decrease overall performance of S. herbacea under earlier snowmelt.  相似文献   

5.
Climate warming is strongly altering the timing of season initiation and season length in the Arctic. Phenological activities are among the most sensitive plant responses to climate change and have important effects at all levels within the ecosystem. We tested the effects of two experimental treatments, extended growing season via snow removal and extended growing season combined with soil warming, on plant phenology in tussock tundra in Alaska from 1995 through 2003. We specifically monitored the responses of eight species, representing four growth forms: (i) graminoids (Carex bigellowii and Eriophorum vaginatum); (ii) evergreen shrubs (Ledum palustre, Cassiope tetragona, and Vaccinium vitis‐idaea); (iii) deciduous shrubs (Betula nana and Salix pulchra); and (iv) forbs (Polygonum bistorta). Our study answered three questions: (i) Do experimental treatments affect the timing of leaf bud break, flowering, and leaf senescence? (ii) Are responses to treatments species‐specific and growth form‐specific? and (iii) Which environmental factors best predict timing of phenophases? Treatment significantly affected the timing of all three phenophases, although the two experimental treatments did not differ from each other. While phenological events began earlier in the experimental plots relative to the controls, duration of phenophases did not increase. The evergreen shrub, Cassiope tetragona, did not respond to either experimental treatment. While the other species did respond to experimental treatments, the total active period for these species did not increase relative to the control. Air temperature was consistently the best predictor of phenology. Our results imply that some evergreen shrubs (i.e., C. tetragona) will not capitalize on earlier favorable growing conditions, putting them at a competitive disadvantage relative to phenotypically plastic deciduous shrubs. Our findings also suggest that an early onset of the growing season as a result of decreased snow cover will not necessarily result in greater tundra productivity.  相似文献   

6.
Many alpine and subalpine plant species exhibit phenological advancements in association with earlier snowmelt. While the phenology of some plant species does not advance beyond a threshold snowmelt date, the prevalence of such threshold phenological responses within plant communities is largely unknown. We therefore examined the shape of flowering phenology responses (linear versus nonlinear) to climate using two long-term datasets from plant communities in snow-dominated environments: Gothic, CO, USA (1974–2011) and Zackenberg, Greenland (1996–2011). For a total of 64 species, we determined whether a linear or nonlinear regression model best explained interannual variation in flowering phenology in response to increasing temperatures and advancing snowmelt dates. The most common nonlinear trend was for species to flower earlier as snowmelt advanced, with either no change or a slower rate of change when snowmelt was early (average 20% of cases). By contrast, some species advanced their flowering at a faster rate over the warmest temperatures relative to cooler temperatures (average 5% of cases). Thus, some species seem to be approaching their limits of phenological change in response to snowmelt but not temperature. Such phenological thresholds could either be a result of minimum springtime photoperiod cues for flowering or a slower rate of adaptive change in flowering time relative to changing climatic conditions.  相似文献   

7.
《植物生态学报》2016,40(10):1028
Aims Climate warming strongly influences reproductive phenology of plants in alpine and arctic ecosystems. Here we focus on phenological shifts caused by warming in a typical alpine meadow on the Qinghai-Xizang Plateau. Our objective was to explore phenological responses of alpine plant species to experimental warming. Methods Passive warming was achieved using open-top chambers (OTCs). The treatments included control (C), and four levels of warming (T1, T2, T3, T4). We selected Kobresia pygmaea, Potentilla saundersiana, Potentilla cuneata, Stipa purpurea, Festuca coelestis and Youngia simulatrix as the focal species. Plant phenology was scored every 3-5 days in the growing season. The reproductive phenology phases of each species were estimated through fitting the phenological scores to the Richards function. Important findings Under soil water stress caused by warming, most plants in the alpine meadow advanced or delayed their reproductive events. As a result, warming significantly delayed phenological development of K. pygmaea. Warming significantly advanced reproductive phenology of P. saundersiana, S. purpurea and F. coelestis, but not of P. cuneata and Y. simulatrix. In addition, warming significantly shortened the average flowering duration of alpine plant species. The potentially warmer and drier growing seasons under climate change may shift the reproductive phenology of the alpine systems in similar pattern.  相似文献   

8.
朱军涛 《植物生态学报》2016,40(10):1028-1036
全球气候变暖对高寒和极地地区的植物物候产生强烈的影响。该研究主要关注增温条件下藏北高寒草甸不同功能型植物繁殖时间(生殖物候)的改变。实验采用开顶箱式增温方法, 对3个主要功能群浅根-早花、浅根-中花和深根-晚花植物的现蕾、开花、结实时间进行观测。研究结果表明: (1)增温导致了土壤水分胁迫, 显著推迟了浅根-早花植物高山嵩草(Kobresia pygmaea)的繁殖时间; (2)增温显著提前了浅根-中花植物钉柱委陵菜(Potentilla saundersiana)和深根晚花植物紫花针茅(Stipa purpurea)和矮羊茅(Festuca coelestis)的繁殖时间; (3)增温没有显著影响浅根-中花植物楔叶委陵菜(Potentilla cuneata)和深根-晚花植物无茎黄鹌菜(Youngia simulatrix)的繁殖时间; (4)增温缩短了3种类型植物的开花持续时间。这些结果显示增温改变了藏北高寒草甸群落中多数物种的繁殖时间, 这预示着在未来更热更干的生长季, 青藏高原高寒草甸系统的植物物候格局可能会被重塑。  相似文献   

9.
The purpose of the experiment was to see whether a change in nutrient availability resulting from perturbations might alter rates of development of the major plant growth forms of an arctic tussock tundra slope. Slow release fertilizer pellets (NPK) were spread in June and July 1985 on the frost surface under an 8-20 cm thick layer of thawed tundra in the northern foothills of the Brooks Range, Alaska. Initiation day of main phenological events, their duration and their intensity (% plants reaching a phenophase in a plot) were recorded during the 1985, 1986 and 1987 growing season. Variations in initiation day of phenophases were mainly due to environmental variations among years and to a lesser degree to fertilization; only timing of vegetative bud break of evergreen shrubs was significantly affected by fertilization, and occurred earlier than in the control. Flowering intensity varied significantly from one year to an other but was not significantly stimulated by fertilization.  相似文献   

10.
Zhu J.-T. 《植物生态学报》2016,(10):1028-1036
Aims: Climate warming strongly influences reproductive phenology of plants in alpine and arctic ecosystems. Here we focus on phenological shifts caused by warming in a typical alpine meadow on the Qinghai-Xizang Plateau. Our objective was to explore phenological responses of alpine plant species to experimental warming. Methods: Passive warming was achieved using open-top chambers (OTCs). The treatments included control (C), and four levels of warming (T1, T2, T3, T4). We selected Kobresia pygmaea, Potentilla saundersiana, Potentilla cuneata, Stipa purpurea, Festuca coelestis and Youngia simulatrix as the focal species. Plant phenology was scored every 3-5 days in the growing season. The reproductive phenology phases of each species were estimated through fitting the phenological scores to the Richards function. Important findings: Under soil water stress caused by warming, most plants in the alpine meadow advanced or delayed their reproductive events. As a result, warming significantly delayed phenological development of K. pygmaea. Warming significantly advanced reproductive phenology of P. saundersiana, S. purpurea and F. coelestis, but not of P. cuneata and Y. simulatrix. In addition, warming significantly shortened the average flowering duration of alpine plant species. The potentially warmer and drier growing seasons under climate change may shift the reproductive phenology of the alpine systems in similar pattern.  相似文献   

11.
Climate change effects on snow cover and thermic regime in alpine tundra might lead to a longer growing season, but could also increase risks to plants from spring frost events. Alpine snowbeds, i.e. alpine tundra from late snowmelt sites, might be particularly susceptible to such climatic changes. Snowbed communities were grown in large monoliths for two consecutive years, under different manipulated snow cover treatments, to test for effects of early (E) and late (L) snowmelt on dominant species growth, plant functional traits, leaf area index (LAI) and aboveground productivity. Spring snow cover was reduced to assess the sensitivity of snowbed alpine species to severe early frost events, and dominant species freezing temperatures were measured. Aboveground biomass, productivity, LAI and dominant species growth did not increase significantly in E compared to L treatments, indicating inability to respond to an extended growing season. Edapho‐climatic conditions could not account for these results, suggesting that developmental constraints are important in controlling snowbed plant growth. Impaired productivity was only detected when harsher and more frequent frost events were experimentally induced by early snowmelt. These conditions exposed plants to spring frosts, reaching temperatures consistent with the estimated freezing points of the dominant species (~?10 °C). We conclude that weak plasticity in phenological response and potential detrimental effects of early frosts explain why alpine tundra from snowbeds is not expected to benefit from increased growing season length.  相似文献   

12.
Under climate warming, plants will undergo novel selective pressures to adjust reproductive timing. Adjustment between reproductive phenology and environment is expected to be higher in arctic and alpine habitats because the growing season is considerably short. As early- and late-flowering species reproduce under very different environmental conditions, selective pressures on flowering phenology and potential effects of climate change are likely to differ between them. However, there is no agreement on the magnitude of the benefits and costs of early- vs. late-flowering species under a global warming scenario. In spite of its relevance, phenotypic selection on flowering phenology has rarely been explored in alpine plants and never in Mediterranean high mountain species, where selective pressures are very different due to the summer drought imposed over the short growth season. We hypothesized that late-flowering plants in Mediterranean mountains should present stronger selective pressures towards early onset of reproduction than early-flowering species, because less water is available in the soil as growing season progresses. We performed selection analyses on flowering onset and duration in two high mountain species of contrasting phenology. Since phenotypic selection can be highly context-dependent, we studied several populations of each species for 2 years, covering their local altitudinal ranges and their different microhabitats. Surrogates of biotic selective agents, like fruitset for pollinators and flower and fruit loss for flower and seed predators, were included in the analysis. Differences between the early- and the late-flowering species were less than expected. A consistent negative correlational selection of flowering onset and duration was found affecting plant fitness, i.e., plants that bloomed earlier flowered for longer periods improving plant fitness. Nevertheless, the late-flowering species may experience higher risks under climate warming because in extremely warm and dry years the earlier season does not bring about a longer flowering duration due to summer drought.  相似文献   

13.
In alpine environments, flowering phenology can differ within local populations even at the same elevation. We assessed the effects of differences in flowering phenology due to snowmelt timing caused by local geographic heterogeneity on the genetic structure of a population of an alpine plant, Erythronium grandiflorum Pursh. We established a study plot of 250×70 m at 3,340 m above sea level in the Front Range of the Rocky Mountains, CO, USA. The flowering phenology was considerably influenced by snowmelt timing due to local geographic heterogeneity. Twenty-two patches of E. grandiflorum were recognized in the study plot and were classified into three phenological groups: early, middle, and late. To express the differentiation of flowering phenology among the patches, we defined phenological distance and analyzed the relationship between genetic and phenological distances. Additionally, since genetic distance is expected to co-vary with geographic distance, we also analyzed the relationship between genetic distance and geographic distance among patches. The results revealed not only that isolation by distance was present among patches, but also that the differences in snowmelt timing gave rise to phenologically distant patches of E. grandiflorum, which in turn determine the genetic structure caused by the limited pollen flow between patches.  相似文献   

14.
Recent changes in climate have led to significant shifts in phenology, with many studies demonstrating advanced phenology in response to warming temperatures. The rate of temperature change is especially high in the Arctic, but this is also where we have relatively little data on phenological changes and the processes driving these changes. In order to understand how Arctic plant species are likely to respond to future changes in climate, we monitored flowering phenology in response to both experimental and ambient warming for four widespread species in two habitat types over 21 years. We additionally used long‐term environmental records to disentangle the effects of temperature increase and changes in snowmelt date on phenological patterns. While flowering occurred earlier in response to experimental warming, plants in unmanipulated plots showed no change or a delay in flowering over the 21‐year period, despite more than 1 °C of ambient warming during that time. This counterintuitive result was likely due to significantly delayed snowmelt over the study period (0.05–0.2 days/yr) due to increased winter snowfall. The timing of snowmelt was a strong driver of flowering phenology for all species – especially for early‐flowering species – while spring temperature was significantly related to flowering time only for later‐flowering species. Despite significantly delayed flowering phenology, the timing of seed maturation showed no significant change over time, suggesting that warmer temperatures may promote more rapid seed development. The results of this study highlight the importance of understanding the specific environmental cues that drive species’ phenological responses as well as the complex interactions between temperature and precipitation when forecasting phenology over the coming decades. As demonstrated here, the effects of altered snowmelt patterns can counter the effects of warmer temperatures, even to the point of generating phenological responses opposite to those predicted by warming alone.  相似文献   

15.
The phenology of arctic ecosystems is driven primarily by abiotic forces, with temperature acting as the main determinant of growing season onset and leaf budburst in the spring. However, while the plant species in arctic ecosystems require differing amounts of accumulated heat for leaf‐out, dynamic vegetation models simulated over regional to global scales typically assume some average leaf‐out for all of the species within an ecosystem. Here, we make use of air temperature records and observations of spring leaf phenology collected across dominant groupings of species (dwarf birch shrubs, willow shrubs, other deciduous shrubs, grasses, sedges, and forbs) in arctic and boreal ecosystems in Alaska. We then parameterize a dynamic vegetation model based on these data for four types of tundra ecosystems (heath tundra, shrub tundra, wet sedge tundra, and tussock tundra), as well as ecotonal boreal white spruce forest, and perform model simulations for the years 1970–2100. Over the course of the model simulations, we found changes in ecosystem composition under this new phenology algorithm compared with simulations with the previous phenology algorithm. These changes were the result of the differential timing of leaf‐out, as well as the ability for the groupings of species to compete for nitrogen and light availability. Regionally, there were differences in the trends of the carbon pools and fluxes between the new phenology algorithm and the previous phenology algorithm, although these differences depended on the future climate scenario. These findings indicate the importance of leaf phenology data collection by species and across the various ecosystem types within the highly heterogeneous Arctic landscape, and that dynamic vegetation models should consider variation in leaf‐out by groupings of species within these ecosystems to make more accurate projections of future plant distributions and carbon cycling in Arctic regions.  相似文献   

16.
Background and Aims Recent global changes, particularly warming and drought, have had worldwide repercussions on the timing of flowering events for many plant species. Phenological shifts have also been reported in alpine environments, where short growing seasons and low temperatures make reproduction particularly challenging, requiring fine-tuning to environmental cues. However, it remains unclear if species from such habitats, with their specific adaptations, harbour the same potential for phenological plasticity as species from less demanding habitats.Methods Fourteen congeneric species pairs originating from mid and high elevation were reciprocally transplanted to common gardens at 1050 and 2000 m a.s.l. that mimic prospective climates and natural field conditions. A drought treatment was implemented to assess the combined effects of temperature and precipitation changes on the onset and duration of reproductive phenophases. A phenotypic plasticity index was calculated to evaluate if mid- and high-elevation species harbour the same potential for plasticity in reproductive phenology.Key Results Transplantations resulted in considerable shifts in reproductive phenology, with highly advanced initiation and shortened phenophases at the lower (and warmer) site for both mid- and high-elevation species. Drought stress amplified these responses and induced even further advances and shortening of phenophases, a response consistent with an ‘escape strategy’. The observed phenological shifts were generally smaller in number of days for high-elevation species and resulted in a smaller phenotypic plasticity index, relative to their mid-elevation congeners.Conclusions While mid- and high-elevation species seem to adequately shift their reproductive phenology to track ongoing climate changes, high-elevation species were less capable of doing so and appeared more genetically constrained to their specific adaptations to an extreme environment (i.e. a short, cold growing season).  相似文献   

17.
Mountain plants are particularly sensitive to climate warming because snowmelt timing exerts a direct control on their reproduction. Current warming is leading to earlier snowmelt dates and longer snow-free periods. Our hypothesis is that high-mountain Mediterranean plants are not able to take advantage of a lengthened snow-free period because this leads to longer drought that truncates the growing season. However, reproductive timing may somewhat mitigate these negative effects through temporal shifts. We assessed the effects of flowering phenology on the reproductive success of Silene ciliata, a Mediterranean high-mountain plant, across an altitudinal gradient during two climatically contrasting years. The species showed a late-flowering pattern hampering the use of snowmelt water. Plant fitness was largely explained by the elapsed time from snowmelt to onset of flowering, suggesting a selective pressure towards early flowering caused by soil moisture depletion. The proportion of flowering plants decreased at the lowest population, especially in the drier year. Plants produced more flowers, fruits and seeds at the highest population and in the mild year. Our results indicate that water deficit in dry years could threaten the lowland populations of this mountainous species, while high-altitude environments are more stable over time.  相似文献   

18.
Changes in growing season temperature and duration may have profound effects on the population dynamics of arctic and alpine plant species in snow-bed and fell-field habitats. We examined how a typical herbaceous pioneer species, Ranunculus glacialis, responded to experimental climate change in open-top chambers for three seasons at an alpine site in southern Norway. Warming had no significant effect on any reproductive, growth or phenological variables, except for seed weight, which increased significantly during the first 2 ears. Despite large differences in average date of snowmelt among years, average reproductive output and ramet size differed little among years. Within-year variation in date of snowmelt had no impact on seed number or weight in either control or warmed plots. Leaf width and ramet leaf number decreased significantly with later snowmelt within a year. Experimental warming reduced the negative effect on ramet size of late snowmelt within a year to some extent. In general, R. glacialis reacts contrary to most other arctic/alpine species to experimental warming. Species with such low responsiveness to environmental conditions may be particularly vulnerable to climatic change, especially if their habitat is invaded by other species with higher phenotypic plasticity and a better competitive ability.  相似文献   

19.
Satellite data indicate significant advancement in alpine spring phenology over decades of climate warming, but corresponding field evidence is scarce. It is also unknown whether this advancement results from an earlier shift of phenological events, or enhancement of plant growth under unchanged phenological pattern. By analyzing a 35‐year dataset of seasonal biomass dynamics of a Tibetan alpine grassland, we show that climate change promoted both earlier phenology and faster growth, without changing annual biomass production. Biomass production increased in spring due to a warming‐induced earlier onset of plant growth, but decreased in autumn due mainly to increased water stress. Plants grew faster but the fast‐growing period shortened during the mid‐growing season. These findings provide the first in situ evidence of long‐term changes in growth patterns in alpine grassland plant communities, and suggest that earlier phenology and faster growth will jointly contribute to plant growth in a warming climate.  相似文献   

20.
Phenological advancement allows individuals to adapt to climate change by timing life‐history events to the availability of key resources so that individual fitness is maximized. However, different trophic levels may respond to changes in their environment at different rates, potentially leading to a phenological mismatch. This may be especially apparent in the highly seasonal arctic environment that is experiencing the effects of climate change more so than any other region. During a 14‐year study near Utqia?vik (formerly Barrow), Alaska, we estimated phenological advancement in egg laying in relation to snowmelt for eight arctic‐breeding shorebirds and investigated potential linkages to species‐specific life‐history characteristics. We found that snowmelt advanced 0.8 days/year—six times faster than the prior 60‐year period. During this same time, six of the eight species exhibited phenological advancement in laying dates (varying among species from 0.1 to 0.9 days earlier per year), although no species appeared capable of keeping pace with advancing snowmelt. Phenological changes were likely the result of high phenotypic plasticity, as all species investigated in this study showed high interannual variability in lay dates. Commonality among species with similar response rates to timing of snowmelt suggests that nesting later and having an opportunistic settlement strategy may increase the adaptability of some species to changing climate conditions. Other life‐history characteristics, such as migration strategy, previous site experience, and mate fidelity did not influence the ability of individuals to advance laying dates. As a failure to advance egg laying is likely to result in greater phenological mismatch, our study provides an initial assessment of the relative risk of species to long‐term climatic changes.  相似文献   

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