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1.
We used experimental transplant studies to understand how dispersal and habitat-specific selection interact to influence plant populations occupying heterogeneous environments. The snow buttercup (Ranunculus adoneus) occupies a steep ecological and flowering time gradient caused by persistent snowmelt differences within its snow bed habitat. We transplanted seeds, seedlings, and adults to learn about the potential interactions between dispersal and selection. We found that adaptive differentiation is not occurring along the snowmelt gradient, despite striking differences in microhabitat conditions and reproductive phenology between early- and latemelting sites. Instead, our results imply that environmentally based differences in seed quality are contributing to directional gene flow from early-melting locations toward latemelting locations. Emergence and early survival of seedlings is greater in late-melting sites in some years, but the larger seeds produced by maternal plants in early-melting locations consistently have a fitness advantage in all parts of the snow bed. Larger seeds survive longer in the soil and have a second peak of seedling emergence in their third year, but these late-emerging seedlings are successful only if dispersed to less vegetated, late-melting destinations. The longer growing season in earlymelting sites enhances vegetative growth at all life-history stages and increases fecundity of seedling transplants but also limits the opportunity for establishment from seed. Our demographic analysis suggests that maternal environmental effects on propagule quality can lead to directional gene flow from benign to marginal sites in populations occupying heterogeneous habitats.  相似文献   

2.
陈文年  吴彦  吴宁  罗鹏 《植物研究》2011,31(2):206-212
在青藏高原东部的一个高山雪床,沿着融雪梯度分别设置早融、中间和晚融3个融雪部位,然后测定川西小黄菊(Pyrethrum tatsienense)、长叶火绒草(Leontopodium longifolium)和圆穗蓼(Polygonum macrophyllum)在3个融雪部位上的物候差异以及种群分布格局的变化。结果表明:从早融到晚融的梯度上,3个物种的物候期都不同程度地有所推迟。其中,开始生长的时间推迟12~14 d,始花期推迟6~8 d,盛花期推迟6 d左右,但同一种植物在不同的融雪部位上的衰老枯黄期趋于一致,这标志着在晚融部位同一植物的生长期要缩短。在种群层次上,长叶火绒草和圆穗蓼的分布格局随着融雪的推迟都发生了一定的变化,基本上表现为从早融部位的集群分布到中间或晚融部位的随机分布。川西小黄菊在各个融雪部位上都表现为集群分布,但集群的强度随融雪的推迟逐渐减弱。  相似文献   

3.
Hirao AS  Kudo G 《Heredity》2004,93(3):290-298
The genetic structure of three snowbed-herb species (Peucedanum multivittatum, Veronica stelleri, and Gentiana nipponica) was analyzed using allozymes across nine populations arranged as a matrix of three snowmelt gradients x three geographic locations within 3 km in the Taisetsu Mountains, northern Japan. Phenologically asynchronous populations are packed within a local area in alpine snowbeds, because flowering season of alpine plants depends strongly on the timing of snowmelt. Moderate genetic differentiation was detected among local populations in every species (FST=0.03-0.07). There was a significant correlation between the geographic distance and genetic distance in the P. multivittatum populations, but not in the V. stelleri and G. nipponica populations. On the other hand, a significant correlation between the phenological distance caused by snowmelt timing and genetic distance was detected in the V. stelleri and G. nipponica populations, but not in the P. multivittatum populations. The snowmelt gradient or geographic separation influenced hierarchical genetic structure of these species moderately (FRT <0.04). Restriction of gene flow due to phenological separation and possible differential selection along the snowmelt gradient may produce genetic clines at microgeographic scale in these species.  相似文献   

4.
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   

5.
This study explores the effects of emergence time and reproductive phenology on seed number, seed size, and seedling survival in a population of the alpine buttercup, Ranunculus adoneus. Phenology in this snow bowl population is structured by snow depth. Plants in late melting interior portions of the bowl emerged and flowered 3 to 4 wk after those in early melting zones at the bowl perimeter during the summers of 1988 and 1989. Flowering time differences of buttercups across the bowl were consistent from one year to the next. In 1988, late flowering plants tended to set fewer seeds than early flowering ones; in 1989 no decrease in seed number accompanied flowering date. Path analysis showed that equal fecundity in early and late emerging portions of the bowl population during 1989 resulted from balancing spatial and temporal constraints on seed production. Spatial aspects of habitat quality improved toward the interior of the bowl, but temporal regimes deteriorated in these late melting sites. In both 1988 and 1989 seed size declined with delays in flowering. Path analysis of 1989 data showed that because of reduced time for seed growth, plants in late melting portions of the bowl set smaller seeds than those in earlier melting zones. Differences in seed size due to parental phenology are likely to influence fitness in snow buttercups. Under natural conditions, seedlings from large seeds (>;0.65 mg) have sixfold higher survival than do those from smaller seeds (<;0.65 mg). We conclude that seedling recruitment may be infrequent in late-melting portions of the snow bowl due to delayed parental phenology.  相似文献   

6.
The timing of the snowmelt is a crucial factor in determining the phenological schedule of alpine plants. A long-term monitoring of snowmelt regimes in a Japanese alpine area revealed that the onset of the snowmelt season has been accelerated during the last 17 years in early snowmelt sites but that such a trend has not been detected in late snowmelt sites. This indicates that the global warming effect on the snowmelt pattern may be site-specific. The flowering phenology of fellfield plants in an exposed wind-blown habitat was consistent between an unusually warm year (1998) and a normal year (2001). In contrast, the flowering occurrence of snowbed plants varied greatly between the years depending on the snowmelt time. There was a large number of flowering species in the fellfield community from mid- to late to late June and from mid- to late July. The flowering peak of an early-melt snowbed plant community was in the middle of the flowering season and that of a late-melt snowbed community was in the early flowering season. These habitat-specific phenological patterns were consistent between 1998 and 2001. The effects of the variation in flowering timing on seed-set success were evaluated for an entomophilous snowbed herb, Peucedanum multivittatum, along the snowmelt gradient during a 5-year period. When flowering occurred prior to early August, mean temperature during the flowering season positively influenced the seed set. When flowering occurred later than early August, however, the plants enjoyed high seed-set success irrespective of temperature conditions if frost damage was absent. These observations are probably explained based on the availability of pollinators, which depends not only on ambient temperature but also on seasonal progress. These results suggest that the effects of climate change on biological interaction may vary depending on the specific habitat in the alpine ecosystem in which diverse snowmelt patterns create complicated seasonality for plants within a very localized area.  相似文献   

7.
Phenological differences in flowering arising along elevational gradients may be caused by either local adaptation or phenotypic plasticity. Local adaptation can lead to reproductive isolation of populations at different elevational zones and thus produce elevational genetic structuring, while phenotypic plasticity does not produce elevational genetic structuring. In this study, we examined the effects of elevation and fragmentation on genetic diversity and structure of Polylepis australis populations, where individuals exhibit phenological differences in flowering along an elevational gradient. We assessed the polymorphism of amplified fragment length polymorphism markers in adults and saplings from one conserved and one fragmented forest covering elevations from 1600 to 2600 m asl. Over 98% of variation was found within populations, and we found very low and similar genetic differentiation along elevational gradients for adults and saplings in both continuous and fragmented forests. In addition, there was no significant relationship between genetic diversity and elevation. Results indicated that phenological differences along elevational gradients are more likely caused by phenotypic plasticity than local adaptation, and fragmentation does not appear to have affected genetic diversity and differentiation in the studied populations. Results therefore imply that if necessary, seeds for reforestation purposes may be collected from different elevations to the seeding or planting sites.  相似文献   

8.
In flowering plants, soil heterogeneity can generate divergent natural selection over fine spatial scales, and thus promote local adaptation in the absence of geographic barriers to gene flow. Here, we investigate phenotypic and genetic differentiation in one of the few flowering plants that thrives in both geothermal and non-thermal soils in Yellowstone National Park (YNP). Yellow monkeyflowers (Mimulus guttatus) growing at two geothermal ("thermal") sites in YNP were distinct in growth form and phenology from paired populations growing nearby (<500?m distant) in non-thermal soils. In simulated thermal and non-thermal environments, thermal plants remained significantly divergent from non-thermal plants in vegetative, floral, mating system, and phenological traits. Plants from both thermal populations flowered closer to the ground, allocated relatively more to sexual reproduction, were more likely to initiate flowering under short daylengths, and made smaller flowers that could efficiently self-fertilize without pollinators. These shared differences are consistent with local adaptation to life in the ephemeral window for growth and reproduction created by winter and spring snowmelt on hot soils. In contrast, habitat type (thermal vs. non-thermal) explained little of the genetic variation at neutral markers. Instead, we found that one thermal population (Agrostis Headquarters; AHQ-T) was strongly differentiated from all other populations (all F (ST)?>?0.34), which were only weakly differentiated from each other (all F (ST)?相似文献   

9.
Current threats to biodiversity, such as climate change, are thought to alter the within-species genetic diversity among microhabitats in highly heterogeneous alpine environments. Assessing the spatial organization and dynamics of genetic diversity within species can help to predict the responses of organisms to environmental change. In this study, we evaluated whether small-scale heterogeneity in snowmelt timing restricts gene flow between microhabitats in the common long-lived dwarf shrub Salix herbacea L. We surveyed 273 genets across 12 early- and late-snowmelt sites (that is, ridges and snowbeds) in the Swiss Alps for phenological variation over 2 years and for genetic variation using seven SSR markers. Phenological differentiation triggered by differences in snowmelt timing did not correlate with genetic differentiation between microhabitats. On the contrary, extensive gene flow appeared to occur between microhabitats and slightly less extensively among adjacent mountains. However, ridges exhibited significantly lower levels of genetic diversity than snowbeds, and patterns of effective population size (Ne) and migration (Nem) between microhabitats were strongly asymmetric, with ridges acting as sources and snowbeds as sinks. As no recent genetic bottlenecks were detected in the studied sites, this asymmetry is likely to reflect current meta-population dynamics of the species dominated by gene flow via seeds rather than ancient re-colonization after the last glacial period. Overall, our results suggest that seed dispersal prevents snowmelt-driven genetic isolation, and snowbeds act as sinks of genetic diversity. We discuss the consequences of such small-scale variation in gene flow and diversity levels for population responses to climate change.  相似文献   

10.
Hirao AS  Kudo G 《Heredity》2008,100(4):424-430
The flowering phenology of alpine-snowbed plants varies widely depending on the time of snowmelt. This variation may cause spatial and temporal heterogeneity in pollen dispersal, which in turn may influence genetic structure. We used spatial autocorrelation analyses to evaluate relative effect of segregation in flowering time and physical distance on fine-scale spatial genetic structure (SGS) of a snowbed herb Primula cuneifolia sampled in 10-m grids within a continuous snow patch (110 x 250 m) using nine allozyme loci. Although the individual flower lasts for 相似文献   

11.
陈文年  吴彦  吴宁  罗鹏  王乾 《生态学报》2011,31(13):3621-3628
在青藏高原东部的的一个高山雪床,沿着融雪梯度设置了早融、中间、晚融三个融雪部位,对每个部位的环境因子、大卫马先蒿的个体生长特征及其繁殖特征进行了测量,并在三个部位间对这些特征进行了比较。无雪期长度、土壤水分含量和表土温度的每日变化幅度在部位间有显著的差异,但土壤营养成份和pH并无明显的变化。从早融部位到晚融部位,大卫马先蒿的株高、单株叶数、单叶面积以及比叶面积显著增加,地上生物产量和总生物量也增加,而地下生物量以及地下生物量与地上生物产量之比却降低。花序中段和下段的花数、单花种子数和种子千粒重随融雪的推迟而增加,花序上段的花数、单花种子数和种子千粒重在融雪梯度上没有明显的变化;就整个花序而言,这些繁殖特征随融雪的推迟而增加。大卫马先蒿的个体生长及繁殖特征主要受冻融交替的影响。  相似文献   

12.
The relationship between flowering phenology and abundance of bumble bees (Bombus spp.) was investigated using 2 years of phenological data collected in an alpine region of northern Japan. Abundance of Bombus species was observed along a fixed transect throughout the flowering season. The number of flowering species was closely related to the floral resources for pollinators at the community scale. In the year with typical weather, the first flowering peak corresponded to the emergence time of queen bees from hibernation, while the second flowering peak corresponded to the active period of worker bees. In the year with an unusually warm spring, however, phenological synchrony between plants and bees was disrupted. Estimated emergence of queen bees was 10 days earlier than the first flowering date owing to earlier soil thawing and warming. However, subsequent worker emergence was delayed, indicating slower colony development. The flowering season finished 2 weeks earlier in the warm-spring year in response to earlier snowmelt. A common resident species in the alpine environment, B. hypocrita sapporoensis, flexibly responded to the yearly fluctuation of flowering. In contrast, population dynamics of other Bombus species were out of synchrony with the flowering: their frequencies were highest at the end of the flowering season in the warm-spring year. Therefore, phenological mismatch between flowers and pollinators is evident during warm years, which may become more prevalent in a warmer climate. To understand the mechanism of phenological mismatch in the pollination system of the alpine ecosystem, ground temperature, snowmelt regime, and life cycle of pollinators are key factors.  相似文献   

13.
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   

14.
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   

15.
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   

16.
Abstract Both genetic differentiation and phenotypic plasticity might be expected to affect the location of geographic range limits. Co‐gradient variation (CoGV), plasticity that is congruent with genetic differentiation, may enhance performance at range margins, whereas its opposite, counter‐gradient variation (CnGV) may hinder performance. Here we report findings of reciprocal transplant experiments intended to tease apart the roles of differentiation and plasticity in producing phenotypic variation across a geographic border between two plant subspecies. Clarkia xantiana ssp. xantiana and C. xantiana ssp. parviflora are California‐endemic annuals that replace each other along a west‐east gradient of declining precipitation. We analyzed variation in floral traits, phenological traits, and vegetative morphological and developmental traits by sowing seeds of 18 populations (six of ssp. xantiana and 12 of ssp. parviflora) at three sites (one in each subspecies' exclusive range and one in the subspecies' contact zone), in two growing seasons (an exceptionally wet El Niño winter and a much drier La Niña winter). Significant genetic differences between subspecies appeared in 11 of 12 traits, and differences were of the same sign as in nature. These findings are consistent with the hypothesis that selection is responsible for subspecies differences. Geographic variation within subspecies over part of the spatial gradient mirrored between‐subspecies differences present at a larger scale. All traits showed significant plasticity in response to spatial and temporal environmental variation. Plasticity patterns ranged from spatial and temporal CoGV (e.g., in node of first flower), to spatial CnGV (e.g., in flowering time), to patterns that were neither CoGV nor CnGV (the majority of traits). Instances of CoGV may reflect adaptive plasticity and may serve to increase performance under year‐to‐year environmental variation and at sites near the subspecies border. However, the presence of spatial CnGV in some critical traits suggests that subspecies ranges may also be constrained by patterns of plasticity.  相似文献   

17.
Reciprocal transplants of both seeds and seedlings were utilized to determine whether populations of the annual grass Amphicarpum purshii have become locally adapted to specific habitats due to the consistent production of cleistogamous subterranean seeds from year to year. The hypothesis was that subterranean seeds placed in the same habitat as the parents will produce seedlings of greater vigor and adults of higher reproductive capacity than plants from seeds transplanted to a different habitat far removed from the parents. For both seed and seedling transplant experiments involving three sites in the Pine Barrens of New Jersey, the effects of site on shoot dry weight and production of aerial spikelets, subterranean spikelets, and seeds were generally much more significant than the effects of population origin. With one exception, there was no tendency for seedlings (or plants from seeds) replanted into their home sites to outperform alien seedlings (or plants from seeds) transplanted into these same sites. The overriding importance of environmental factors (relative to genetic differences among populations) in determining the phenotypic expression of life history characters, and selection occurring during succession at a site may retard the evolution of genetic adaptation to local habitat conditions in this species.  相似文献   

18.
Flowering phenology of alpine plants is strongly determined by the timing of snowmelt, and the conditions of pollination of widely distributed plants vary greatly during their flowering season. We examined the reproductive success of the distylous alpine herb, Primula modesta, along the snowmelt gradient under natural conditions, and compared it with the result of artificial pollination experiments. In addition, the compositions and visit frequencies of pollinators to the flower of P. modesta were examined during the flowering period. The pin and thrum plants of P. modesta growing at the same site have an equal ability to produce seeds if a sufficient amount of legitimate pollen grains are deposited on the stigma surface. However, under natural conditions, their seed‐set success was often (even if not always) restricted by pollen limitation, and the functional gender of the pin and thrum plants biased to the female and male, respectively, associated with their growing sites. These variations were not ascribed to resource limitation nor biased morph ratio but to the seasonal changes in pollination situations, a replacement of pollinator types from long‐ to short‐tongued pollinators resulted in unidirectional pollen transfer from long stamens (thrum plants) to long styles (pin plants). The functional gender specialization may enhance the evolution of dioecy from heterostyly, but the severe pollen limitation may cause the breakdown of heterostyly into homostyly. To consider the evolutionary pathway of heterostylous plants, an accumulation of the empirical data is required demonstrating how phenological synchrony between plants and pollinators is decided and to what degree this relationship is stable over years, along with estimates of selection and gene flow in individual plants.  相似文献   

19.

Background

Quercus oleoides Cham. and Schlect., tropical live oak, is a species of conservation importance in its southern range limit of northwestern Costa Rica. It occurs in high-density stands across a fragmented landscape spanning a contrasting elevation and precipitation gradient. We examined genetic diversity and spatial genetic structure in this geographically isolated and genetically distinct population. We characterized population genetic diversity at 11 nuclear microsatellite loci in 260 individuals from 13 sites. We monitored flowering time at 10 sites, and characterized the local environment in order to compare observed spatial genetic structure to hypotheses of isolation-by-distance and isolation-by-environment. Finally, we quantified pollen dispersal distances and tested for local adaptation through a reciprocal transplant experiment in order to experimentally address these hypotheses.

Results

High genetic diversity is maintained in the population and the genetic variation is significantly structured among sampled sites. We identified 5 distinct genetic clusters and average pollen dispersal predominately occurred over short distances. Differences among sites in flowering phenology and environmental factors, however, were not strictly associated with genetic differentiation. Growth and survival of upland and lowland progeny in their native and foreign environments was expected to exhibit evidence of local adaptation due to the more extreme dry season in the lowlands. Seedlings planted in the lowland garden experienced much higher mortality than seedlings in the upland garden, but we did not identify evidence for local adaptation.

Conclusion

Overall, this study indicates that the Costa Rican Q. oleoides population has a rich population genetic history. Despite environmental heterogeneity and habitat fragmentation, isolation-by-distance and isolation-by-environment alone do not explain spatial genetic structure. These results add to studies of genetic structure by examining a common, tropical tree over multiple habitats and provide information for managers of a successional forest in a protected area.  相似文献   

20.
The idiosyncratic behaviours of seed dispersers are important contributors to plant spatial associations and genetic structures. In this study, we used a combination of field, molecular and spatial studies to examine the connections between seed dispersal and the spatial and genetic structures of a dominant neotropical palm Attalea phalerata. Field observation and genetic parentage analysis both indicated that the majority of A. phalerata seeds were dispersed locally over short distances (<30 m from the maternal tree). Spatial and genetic structures between adults and seedlings were consistent with localized and short-distance seed dispersal. Dispersal contributed to spatial associations among maternal sibling seedlings and strong spatial and genetic structures in both seedlings dispersed near (<10 m) and away (>10 m) from maternal palms. Seedlings were also spatially aggregated with juveniles. These patterns are probably associated with the dispersal of seeds by rodents and the survival of recruits at specific microsites or neighbourhoods over successive fruiting periods. Our cross-cohort analyses found palms in older cohorts and cohort pairs were associated with a lower proportion of offspring and sibling neighbours and exhibited weaker spatial and genetic structures. Such patterns are consistent with increased distance- and density-dependent mortality over time among palms dispersed near maternal palms or siblings. The integrative approaches used for this study allowed us to infer the importance of seed dispersal activities in maintaining the aggregated distribution and significant genetic structures among A. phalerata palms. We further conclude that distance- and density-dependent mortality is a key postdispersal process regulating this palm population.  相似文献   

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