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1.
Animals exhibit varied life‐history traits that reflect adaptive responses to their environments. For Arctic‐breeding birds, traits related to diet, egg nutrient allocation, clutch size, and chick growth are predicted to be under increasing selection pressure due to rapid climate change and increasing environmental variability across high‐latitude regions. We compared four migratory birds (black brant [Branta bernicla nigricans], lesser snow geese [Chen caerulescens caerulescens], semipalmated sandpipers [Calidris pusilla], and Lapland longspurs [Calcarius lapponicus]) with varied life histories at an Arctic site in Alaska, USA, to understand how life‐history traits help moderate environmental variability across different phases of the reproductive cycle. We monitored aspects of reproductive performance related to the timing of breeding, reproductive investment, and chick growth from 2011 to 2018. In response to early snowmelt and warm temperatures, semipalmated sandpipers advanced their site arrival and bred in higher numbers, while brant and snow geese increased clutch sizes; all four species advanced their nest initiation dates. During chick rearing, longspur nestlings were relatively resilient to environmental variation, whereas warmer temperatures increased the growth rates of sandpiper chicks but reduced growth rates of snow goose goslings. These responses generally aligned with traits along the capital‐income spectrum of nutrient acquisition and altricial–precocial modes of chick growth. Under a warming climate, the ability to mobilize endogenous reserves likely provides geese with relative flexibility to adjust the timing of breeding and the size of clutches. Higher temperatures, however, may negatively affect the quality of herbaceous foods and slow gosling growth. Species may possess traits that are beneficial during one phase of the reproductive cycle and others that may be detrimental at another phase, uneven responses that may be amplified with future climate warming. These results underscore the need to consider multiple phases of the reproductive cycle when assessing the effects of environmental variability on Arctic‐breeding birds.  相似文献   

2.
Plants from four populations of Hordeum spontaneum originating in distinct environments of Israel were compared for stress induced phenotypic plasticity. The environments ranged along a gradient of increasing rainfall amount and predictability from low (desert) to moderate (semisteppe batha) to high (Mediterranean grassland and mountain, the latter also experiencing frost stress). The plants were exposed to a set of four treatments: no stress (optimum water and nutrients), water, nutrient and both water and nutrient stress. Plants from the four populations (or ecotypes) exhibited different patterns of plasticity in response to the different stresses (water and nutrients) and in different trait categories (reproductive, fitness and resource allocation). The importance of plasticity in response to water stress appears to decrease, and to nutrient stress appears to increase along the increasing rainfall gradient. The mountain ecotype, growing in an area with high potential productivity (amount of rainfall) but experiencing periodic frosts, was the most plastic among ecotypes in resource allocation under both water and nutrient stress, but exhibited low plasticity in other trait categories. In contrast, the desert ecotype had low plasticity in resource allocation under water stress and the lowest plasticity among the four ecotypes in all trait categories in response to nutrient stress. The ecotype originating in Mediterranean grassland, a predictable and most favourable environment, was highly plastic in fitness and allocation traits in response to low nutrient levels which is likely to occur due to competition in productive environment. We discuss the observed differences in ecotype plasticity as part of their environmentally induced adaptive ‘strategies’. We found no support for the hypothesis that plants originating in environments with greater variation and unpredictability are more plastic. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society 2002, 75 , 301–312.  相似文献   

3.
Substantial inter- and intraspecific variation is found in reproductive traits, but the evolutionary implications of this variation remain unclear. One hypothesis is that natural selection favours female reproductive morphology that allows females to control mating and fertilization and that diverse male reproductive traits arise as counter adaptations to subvert this control. Such co-evolution predicts the establishment of genetic correlations between male and female reproductive traits that closely interact during mating. Therefore, we measured phenotypic and genetic correlations between male and female reproductive tract characteristics in the yellow dung fly, Scathophaga stercoraria (Diptera: Scathophagidae), using a nested half-sib breeding experiment. We found significant heritabilities for the size of most reproductive tract traits investigated in both females (spermathecae and their ducts, accessory glands and their ducts) and males (testis size but not sperm length). Within the sexes, phenotypic and genetic correlations were mostly nil or positive, suggesting functional integration of or condition-dependent investment in internal reproductive traits. Negative intrasexual genetic correlations, potentially suggestive of resource allocation trade-offs, were not evident. Intersexual genetic correlations were mostly positive, reflecting expected allometries between male and female morphologies. Most interestingly, testis size correlated positively with female accessory gland size and duct length, potentially indicative of a co-evolutionary arms race. We discuss these and alternative explanations for these patterns of genetic covariance.  相似文献   

4.
5.
The spawning period of the common goby Pomatoschistus microps from 1993 to 1997 in the Vaccarès lagoon did not vary, except in 1997 when it was longer due to the reproduction of the young-of-the-year. Egg size and number, and reproductive allocation varied greatly with one year to another. Female common gobies increased both their fecundity per spawning act and their egg size from 1993 to 1995. The annual variation in the reproductive effort suggests a high phenotypic plasticity of reproductive traits in P. microps , in the face of environmental perturbations. In winter 1993–1994, a centennial flood of the Rhône River caused major hydrological changes in the lagoon in less than 1 week, affecting many invertebrates and fish for several years. The reproductive investment of the common goby increased, possibly as a consequence of those environmental changes.  相似文献   

6.
研究植物群落功能性状间的相关关系及其对环境变化的响应,能够有效揭示植物功能性状的权衡模式及其对环境的适应策略。藏东昌都地区位于横断山脉西北部,复杂气候地貌孕育了丰富的植物资源,是青藏高原森林灌丛生态系统主要组分和国际生物多样性保护的热点地区。以藏东森林灌丛群落优势木本植物为研究对象,在大量野外调查基础上,采用相关分析、主成分分析、线性回归和方差分析等方法,研究了该区域植物功能性状间的相关关系、功能性状对环境变量的响应规律以及功能性状的变异来源。结果表明:(1)藏东木本植物表现出适应高寒环境的性状权衡模式,即:比叶面积、叶体积较小而叶干物质含量较大,叶磷含量和叶钾含量协同变化;(2)海拔和气候变量共同驱动着藏东木本植物功能性状的变化,并且藏东木本植物倾向于采取“高投入—慢回报”提高御寒能力的保守型适应策略;(3)海拔是影响藏东植物功能性状变异最显著的环境变量,种间变异在藏东植物群落功能性状随环境变化中起主要作用。研究结果揭示了藏东木本植物功能性状的权衡模式及其对高寒环境的适应策略,有助于加深对藏东自然植物资源分布规律和生态功能的认识,为区域生态系统功能和生物多样性保护提供科学依据。  相似文献   

7.
Evidence synthesis, both qualitatively and quantitatively through meta-analysis, is central to the development of evidence-based medicine. Unfortunately, meta-analysis is often complicated by the suspicion that the available studies represent a biased subset of the evidence, possibly due to publication bias or other systematically different effects in small studies. A number of statistical methods have been proposed to address this, among which the trim-and-fill method and the Copas selection model are two of the most widely discussed. However, both methods have drawbacks: the trim-and-fill method is based on strong assumptions about the symmetry of the funnel plot; the Copas selection model is less accessible to systematic reviewers, and sometimes encounters estimation problems. In this article, we adopt a logistic selection model, and show how treatment effects can be rapidly estimated via multiple imputation. Specifically, we impute studies under a missing at random assumption, and then reweight to obtain estimates under nonrandom selection. Our proposal is computationally straightforward. It allows users to increase selection while monitoring the extent of remaining funnel plot asymmetry, and also visualize the results using the funnel plot. We illustrate our approach using a small meta-analysis of benign prostatic hyperplasia.  相似文献   

8.
In the wet-dry tropics of northern Australia, temperatures are high and stable year-round but monsoonal rainfall is highly seasonal and variable both annually and spatially. Many features of reproduction in vertebrates of this region may be adaptations to dealing with this unpredictable variation in precipitation, notably by (i) using direct proximate (rainfall-affected) cues to synchronize the timing and extent of breeding with rainfall events, (ii) placing the eggs or offspring in conditions where they will be buffered from rainfall extremes, and (iii) evolving developmental plasticity, such that the timing and trajectory of embryonic differentiation flexibly respond to local conditions. For example, organisms as diverse as snakes (Liasis fuscus, Acrochordus arafurae), crocodiles (Crocodylus porosus), birds (Anseranas semipalmata) and wallabies (Macropus agilis) show extreme annual variation in reproductive rates, linked to stochastic variation in wet season rainfall. The seasonal timing of initiation and cessation of breeding in snakes (Tropidonophis mairii) and rats (Rattus colletti) also varies among years, depending upon precipitation. An alternative adaptive route is to buffer the effects of rainfall variability on offspring by parental care (including viviparity) or by judicious selection of nest sites in oviparous taxa without parental care. A third type of adaptive response involves flexible embryonic responses (including embryonic diapause, facultative hatching and temperature-dependent sex determination) to incubation conditions, as seen in squamates, crocodilians and turtles. Such flexibility fine-tunes developmental rates and trajectories to conditions--especially, rainfall patterns--that are not predictable at the time of oviposition.  相似文献   

9.
Under global warming, the survival of many populations of sedentary organisms in seasonal environments will largely depend on their ability to cope with warming in situ by means of phenotypic plasticity or adaptive evolution. This is particularly true in high‐latitude environments, where current growing seasons are short, and expected temperature increases large. In such short‐growing season environments, the timing of growth and reproduction is critical to survival. Here, we use the unique setting provided by a natural geothermal soil warming gradient (Hengill geothermal area, Iceland) to study the response of Cerastium fontanum flowering phenology to temperature. We hypothesized that trait expression and phenotypic selection on flowering phenology are related to soil temperature, and tested the hypothesis that temperature‐driven differences in selection on phenology have resulted in genetic differentiation using a common garden experiment. In the field, phenology was related to soil temperature, with plants in warmer microsites flowering earlier than plants at colder microsites. In the common garden, plants responded to spring warming in a counter‐gradient fashion; plants originating from warmer microsites flowered relatively later than those originating from colder microsites. A likely explanation for this pattern is that plants from colder microsites have been selected to compensate for the shorter growing season by starting development at lower temperatures. However, in our study we did not find evidence of variation in phenotypic selection on phenology in relation to temperature, but selection consistently favoured early flowering. Our results show that soil temperature influences trait expression and suggest the existence of genetically based variation in flowering phenology leading to counter‐gradient local adaptation along a gradient of soil temperatures. An important implication of our results is that observed phenotypic responses of phenology to global warming might often be a combination of short‐term plastic responses and long‐term evolutionary responses, acting in different directions.  相似文献   

10.
Plant persistence and migration in face of climate change depends on successful reproduction by seed, a central aspect of plant life that drives population dynamics, community assembly and species distributions. Plant reproduction by seed is a chain of physiological processes, the rates of which are a function of temperature, and can be modelled using thermal time models. Importantly, while seed reproduction responds to its instantaneous thermal environment, there is also evidence of phenotypic plasticity in response to the thermal history experienced by the plant's recent ancestors, by the reproducing plant since seedling establishment, and by its seeds both before and after their release. This phenotypic plasticity enables a thermal memory of plant reproduction, which allows individuals to acclimatise to their surroundings. This review synthesises current knowledge on the thermal memory of plant reproduction by seed, and highlights its importance for modelling approaches based on physiological thermal time. We performed a comprehensive search in the Web of Science and analysed 533 relevant articles, of which 81 provided material for a meta‐analysis of thermal memory in reproductive functional traits based on the effect size Zr. The articles encompassed the topics of seed development, seed yield (mass and number), seed dormancy (physiological, morphological and physical), germination, and seedling establishment. The results of the meta‐analysis provide evidence for a thermal memory of seed yield, physiological dormancy and germination. Seed mass and physiological dormancy appear to be the central hubs of this memory. We argue for integrating thermal memory into a predictive framework based on physiological time modelling. This will provide a quantitative assessment of plant reproduction, a complex system that integrates past and present thermal inputs to achieve successful reproduction in changing environments. The effects of a warming environment on plant reproduction cannot be reduced to a qualitative interpretation of absolute positives and negatives. Rather, these effects need to be understood in terms of changing rates and thresholds for the physiological process that underlie reproduction by seed.  相似文献   

11.
A trade‐off between reproduction and somatic maintenance and hence survival is fundamental to life‐history theory. We investigated the relationship between female fecundity and longevity in Homo sapiens using data from 153 countries located all over the world. The raw correlation between life span and fecundity was highly significant with a negative trend. After longevity and fecundity estimates were controlled for by confounding factors such as historical (i.e. human ethnic groups), religious, geographical, socio‐economical and parasitological components, we still observed a negative relationship between the mean female fecundity and the mean longevity in a country. These findings support the hypothesis for the existence of a trade‐off between these two key life‐history traits in humans, as also reported by a recent single longitudinal study in England.  相似文献   

12.
通过2006年11月至2008年8月2周年8个季度的野外采样调查和室内分析,研究了新疆乌伦古湖浮游甲壳动物的季节演替格局及其与环境因子的相关性。结果表明,枝角类和桡足类的总种类数为25种,其中枝角类6种,桡足类19种。枝角类Shannon-Wiener指数2007年夏季和2008年夏季较高,2007年冬季最低;Pielou 指数2006年冬季最高;桡足类Shannon-wiener指数和Pielou指数除冬季较低外,其余三季较高。方差分析结果表明枝角类和无节幼体的密度与生物量呈现出显著的季节变动格局,其中,枝角类密度和生物量在夏秋季形成单一高峰,最大值出现在2007年夏,最大密度和生物量分别为2.12个/L和0.062 mg/L,而无节幼体密度和生物量形成春秋双高峰,其最大值出现在2007年春,密度和生物量分别为5.14个/L和0.015 mg/L,桡足类密度和生物量季节变化不明显。优势种的季节演替方面,象鼻溞和模式有爪猛水蚤全年形成优势,其他枝角类、哲水蚤和剑水蚤不同季节的优势种各异。经典范对应分析(CCA)结果表明,长刺溞等与水体中的pH值、SD呈正相关,与浮游植物生物量呈负相关;长额象鼻溞等与水深呈正相关,无节幼体等与水温也呈正相关性。  相似文献   

13.
根据2011年8-9月南沙群岛海域的生物和环境调查资料,对网采浮游植物的群落特征及其与环境因子的关系进行分析.结果表明: 在鉴定的3门34属113种网采浮游植物中,甲藻门和硅藻门种类各占57.5%和40.7%,其中硅藻门的角藻属种类最多,占30.1%.网采浮游植物平均丰度为2.12×104 cell·m-3,丰度高值区出现在北部礼乐滩多涡区域和南部万安气旋涡附近海域.蓝藻门束毛藻属是网采浮游植物的主要功能群,占海域平均丰度的77.0%.主要优势种为铁氏束毛藻、红海束毛藻和夜光梨甲藻.优势种(类)存在较明显的空间差异,蓝藻门在S3、S5、S6和S10~S14站占优势;甲藻门在中部S4、S7~S9站占优势,硅藻门在南部S1和S2站占优势.网采浮游植物多样性和均匀度指数分别为3.10和0.62.影响浮游植物群落特征的重要环境因子有盐度、水温、铵氮、亚硝酸盐、磷酸盐和硅酸盐,以及南沙中尺度气旋涡和南沙西部沿岸流.典范对应分析排序图较好地显示了浮游植物和环境因子之间的关系.  相似文献   

14.
李晓玲  温浩然  王雪松  杨进  黄成名 《生态学报》2018,38(10):3581-3591
探究中华蚊母树在不同生境水文节律下的生态适应性——表型可塑性,以及它的土壤环境影响因子,对其在异质生境下的生存适应策略具有重要的生态学意义。调查中华蚊母树在自然消落带、反季节消落带及无消涨节律生长区等3种异质生境下36个不同种群叶片主要结构性状叶长(LL)、叶宽(LW)、叶面积(LA)、叶干物重(LDW)以及比叶面积(SLA)的可塑性及其土壤环境影响因子。对各异质生境下中华蚊母树叶片性状的差异显著性进行分析,将LL、LW、LA、LDW之间的异速生长关系用经典异速方程来描述。运用典范对应分析(CCA)研究中华蚊母树叶片性状与土壤环境因子之间的关系。结果显示,中华蚊母树种群的LL、LW、LA、LDW及SLA等5个叶片功能性状在异质生境之间均有显著性差异(P0.05),变异系数平均值在4.80%—26.12%之间,其中LA和SLA在各生境中变异系数最大。各异质生境下中华蚊母树LW与LL、LDW与LL、LW及LA均呈显著的幂函数异速生长关系。其异速系数lgβ均表现极显著差异(P0.01),但其异速指数ɑ却表现相同的规律:LW与LLɑ1,LDW与LL、LWɑ1,LDW与LAɑ≈1,即在不同生境下叶片各性状的生长速率均表现为:LA≈LDWLLLW。各异质生境中叶片较大的表型可塑性和一致的异速生长规律表明了中华蚊母树面对不同的水文节律具有较宽的生态幅,且表现出生长的一致性。CCA排序结果显示,全磷(TP)、水解氮(AN)、速效钾(AK)及土壤含水量(SWC)是中华蚊母树叶片表型可塑性的主要影响因素,中华蚊母树主要通过这些土壤环境因子的驱动改变叶片结构性状使其能在不同水文节律的异质生境中表现出最佳的适合度。  相似文献   

15.
Conditional strategies are the most common form of discrete phenotypic plasticity. In a conditional strategy, the phenotype expressed by an organism is determined by the difference between an environmental cue and a threshold, both of which may vary among individuals. The environmental threshold model (ETM) has been proposed as a mean to understand the evolution of conditional strategies, but has been surprisingly seldom applied to empirical studies. A hindrance for the application of the ETM is that often, the proximate cue triggering the phenotypic expression and the individual threshold are not measurable, and can only be assessed using a related observable cue. We describe a new statistical model that can be applied in this common situation. The Latent ETM (LETM) allows for a measurement error in the phenotypic expression of the individual environmental cue and a purely genetically determined threshold. We show that coupling our model with quantitative genetic methods allows an evolutionary approach including an estimation of the heritability of conditional strategies. We evaluate the performance of the LETM with a simulation study and illustrate its utility by applying it to empirical data on the size-dependent smolting process for stream-dwelling Atlantic salmon juveniles.  相似文献   

16.
Rapidly rising temperatures are expected to cause latitudinal and elevational range shifts as species track their optimal climate north and upward. However, a lack of adaptation to environmental conditions other than climate – for example photoperiod, biotic interactions, or edaphic conditions – might limit the success of immigrants in a new location despite hospitable climatic conditions. Here, we present one of the first direct experimental tests of the hypothesis that warmer temperatures at northern latitudes will confer a fitness advantage to southern immigrants relative to native populations. As rates of warming in the Arctic are more than double the global average, understanding the impacts of warming in Arctic ecosystems is especially urgent. We established experimentally warmed and nonwarmed common garden plots at Alexandra Fiord, Ellesmere Island in the Canadian High Arctic with seeds of two forb species (Oxyria digyna and Papaver radicatum) originating from three to five populations at different latitudes across the Arctic. We found that plants from the local populations generally had higher survival and obtained a greater maximum size than foreign individuals, regardless of warming treatment. Phenological traits varied with latitude of the source population, such that southern populations demonstrated substantially delayed leaf‐out and senescence relative to northern populations. Our results suggest that environmental conditions other than temperature may influence the ability of foreign populations and species to establish at more northerly latitudes as the climate warms, potentially leading to lags in northward range shifts for some species.  相似文献   

17.
Animals make behavioural and reproductive decisions that maximise their lifetime reproductive success, and thus their fitness, in light of periodic and stochastic variability of the environment. Modelling the variation of an individual's energy levels formalises this tradeoff and helps to quantify the population‐level consequences of stressors (e.g. disturbance from human activities and environmental change) that can affect behaviour or physiology. In this study, we develop a dynamic state variable model for the spatially explicit behaviour, physiology and reproduction of a female, long‐lived, migratory marine vertebrate. The model can be used to investigate the spatio‐temporal patterns of behaviour and reproduction that allow an individual to maximise its overall reproductive output. We parametrised the model for eastern North Pacific blue whales Balaenoptera musculus, and used it to predict the effects of changing environmental conditions and increasing human disturbance on the population's vital rates. In baseline conditions, the model output had high fidelity to observed energy dynamics, movement patterns and reproductive strategies. Simulated scenarios suggested that environmental changes could have severe consequences on the population's vital rates, but that individuals could tolerate high levels of anthropogenic disturbance. However, this ability depended on where, when and how often disturbance occurred. In scenarios with both environmental change and anthropogenic disturbance, synergistic interactions caused stronger effects than in isolation. In general, larger body size offered a buffer against stochasticity and disturbance, and, consequently, we predicted juveniles to be more susceptible to disturbance. We also predicted that females prioritise their own survival at the expense of the current reproductive attempt, presumably the result of their long lifespan. Our approach provides a general framework to make predictions of the cumulative and synergistic effects of human disturbance and climate change on migratory populations, which can inform effective management and conservation efforts.  相似文献   

18.
Predicting the response of species to environmental changes is a great and on‐going challenge for ecologists, and this requires a more in‐depth understanding of the importance of biotic interactions and the population structuration in the landscape. Using a reciprocal transplantation experiment, we tested the response of five species to an elevational gradient. This was combined to a neighbour removal treatment to test the importance of local adaptation and biotic interactions. The trait studied was performance measured as survival and biomass. Species response varied along the elevational gradient, but with no consistent pattern. Performance of species was influenced by environmental conditions occurring locally at each site, as well as by positive or negative effects of the surrounding vegetation. Indeed, we observed a shift from competition for biomass to facilitation for survival as a response to the increase in environmental stress occurring in the different sites. Unlike previous studies pointing out an increase of stress along the elevation gradient, our results supported a stress gradient related to water availability, which was not strictly parallel to the elevational gradient. For three of our species, we observed a greater biomass production for the population coming from the site where the species was dominant (central population) compared to population sampled at the limit of the distribution (marginal population). Nevertheless, we did not observe any pattern of local adaptation that could indicate adaptation of populations to a particular habitat. Altogether, our results highlighted the great ability of plant species to cope with environmental changes, with no local adaptation and great variability in response to local conditions. Our study confirms the importance of taking into account biotic interactions and population structure occurring at local scale in the prediction of communities’ responses to global environmental changes.  相似文献   

19.
Abstract. Question: How do the relative frequencies of plant traits (clonality, growth form, seed weight, diaspore morphology) vary during the life cycle and how does this affect regeneration? Location: Alpine meadow and heath communities at Kilpisjärvi, sub‐Arctic Finland. Methods: Control plots and three treatments were used to measure relative species abundances for five life cycle stages: standing vegetation, seed rain, seed bank and seedlings emerging in gaps and in closed vegetation. Results: The relative frequencies of plant traits varied between the life cycle stages. The meadows were dominated by weakly clonal herbs, small or intermediate seeds and unappendaged diaspores, while the heaths were dominated by clonal dwarf shrubs, small seeds and fleshy fruits. In the meadows, species with small seeds dominated during the seed rain and in the seedling stage in gaps, while species with intermediate seeds dominated the seed bank and the seedling stage in closed vegetation. Species with unappendaged diaspores dominated throughout the life cycle. In the heaths, seed bank and seedling stage were practically absent. Conclusions: The observed differences in plant trait spectra between life cycle stages indicate that important environmental factors differ among the stages. Small seeds are advantageous for dispersal, whereas intermediate seeds have a greater probability of germinating and establishing in closed vegetation. Appendages facilitate dispersal, whereas unappendaged diaspores favour seed burial. Although the plant growth form spectrum largely reflects environmental constraints during the regeneration cycle, information on seed weight and diaspore morphology improves our knowledge of the relative importance of morphological adaptations of sexual structures in different stages during the life cycle.  相似文献   

20.
Summary 1  Associations between abiotic variables and patterns of species distribution and abundance are a major preoccupation of community ecologists. In many habitats, this association is manifest in discrete zones of vegetation.
2  We used statistical methods to examine tree species distribution patterns in relatively undisturbed regions of the Sundarbans of Bangladesh. We tested the hypothesis that mangroves occur in discrete zones with respect to elevation. These data were gathered with explicit attention to local and regional differences in edaphic characteristics so that species-environment relationships could be analysed at several spatial scales.
3  Correlations were also assessed between mangrove species composition and edaphic variables that co-vary with elevation, i.e. salinity, field capacity, cation exchange capacity, percentage silt, and mangrove physiognomic category (slope, basin, levee and flat).
4  Quantitative statistical analysis using randomization techniques failed to detect species zonation along any of 33 individual 200-m transects, within 1-km2 blocks, or within 1200-km2 regions.
5  Canonical correspondence analysis relating edaphic variables to species distributions accounted for a total of only 24% of the variance in species composition.
6  Our data suggest that the absence of zonation in the Bangladesh Sundarbans reflects the underlying biology of the system and is not an artefact of long-term human disturbance.  相似文献   

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