Biodiversity, distributions and adaptations of Arctic species in the context of environmental change

The individual of a species is the basic unit which responds to climate and UV-B changes, and it responds over a wide range of time scales. The diversity of animal, plant and microbial species appears to be low in the Arctic, and decreases from the boreal forests to the polar deserts of the extreme North but primitive species are particularly abundant. This latitudinal decline is associated with an increase in super-dominant species that occupy a wide range of habitats. Climate warming is expected to reduce the abundance and restrict the ranges of such species and to affect species at their northern range boundaries more than in the South: some Arctic animal and plant specialists could face extinction. Species most likely to expand into tundra are boreal species that currently exist as outlier populations in the Arctic. Many plant species have characteristics that allow them to survive short snow-free growing seasons, low solar angles, permafrost and low soil temperatures, low nutrient availability and physical disturbance. Many of these characteristics are likely to limit species' responses to climate warming, but mainly because of poor competitive ability compared with potential immigrant species. Terrestrial Arctic animals possess many adaptations that enable them to persist under a wide range of temperatures in the Arctic. Many escape unfavorable weather and resource shortage by winter dormancy or by migration. The biotic environment of Arctic animal species is relatively simple with few enemies, competitors, diseases, parasites and available food resources. Terrestrial Arctic animals are likely to be most vulnerable to warmer and drier summers, climatic changes that interfere with migration routes and staging areas, altered snow conditions and freeze-thaw cycles in winter, climate-induced disruption of the seasonal timing of reproduction and development, and influx of new competitors, predators, parasites and diseases. Arctic microorganisms are also well adapted to the Arctic's climate: some can metabolize at temperatures down to -39 degrees C. Cyanobacteria and algae have a wide range of adaptive strategies that allow them to avoid, or at least minimize UV injury. Microorganisms can tolerate most environmental conditions and they have short generation times which can facilitate rapid adaptation to new environments. In contrast, Arctic plant and animal species are very likely to change their distributions rather than evolve significantly in response to warming.     

人为因素对社会-生态系统脆弱性和恢复力影响的地理差异

横跨北极的地区,可更新资源和不可更新资源、人口密度、资产投资、住宅与交通基础设施的分布存在着很大差异,这些差异被公认为是研究当前和未来社会-生态系统变化的热点.由于当前政治和经济的稳定,芬诺斯堪的亚地区北部社会-生态系统呈现出相对良好的形势,而俄罗斯北部从苏联解体开始便一直经历着急速且消极的变化,这些变化反映了俄罗斯普遍存在的危机状态.北美洲虽然处在一个相对稳定有序的制度下,采取了一些措施用于防止和减轻与工业有关的环境退化,但大多即将出台的新发展计划仍将会显著扩大社会-生态系统受到潜在影响的空间范围.制度或管理条款与生态系统服务功能对环境变化的缓冲能力紧密相关.无论气候是否变暖,某些地理区域对外来破坏作用表现得特别脆弱,这有可能威胁到这些区域在将来提供产品和服务的能力.尽管某些地区的气候变化有可能会使这种形势恶化,但从长远来看,气候变化也有可能会使其恢复力得到加强.     

北极陆地生态系统、气候及紫外线辐射的历史演变

在末次盛冰期,地球上很多大陆都被大量的冰层所覆盖,一些浅海域的海床露出水面将先前分离的大陆连接起来.尽管存在一些适宜于动植物生长而未被冰层覆盖的地区,但其年平均气温仍比更新世时期低10～13℃.在盏冰期的几千年时间内冰川开始消融,其显著特征是气候在大约18000～11400年以前出现了一系列的波动.气候在更新世度过一个温暖期后,开始了一个缓慢的全面变冷的过程,这导致了一系列为期几百年至几千年的气候波动,例如发生在大约13世纪晚期至19世纪早期的"小冰期".在最近150000年的气候变化过程中,北极的各种生态系统和生物组成在近10000年接近其最低分布范围.大冰期结束时的全球大范围急剧升温导致了许多物种的消失,这使北极地区的生物多样性大大降低.因此,北极生态系统以及大型脊椎动物等北极生物的生存正在受到威胁,尤其是目前以及将来的全球变暖都会进一步给它们带来重大灾难.已有的证据表明,就像更新世早期的情形那样,北极地区的树线很有可能会进一步向北发展,并迅速进入到苔原地区,从而减小苔原带,这就会进一步增加北极地区物种灭绝的可能性.一些物种将很有可能向北扩大它们的领地,并取代该地区原有的物种.在更新世早期,由于北极地区的海平面相对较低,当树线入侵到现在的海岸地区时,苔原带至少能够在北极圈的一部分低地区域生存,而从目前来看,未来的海平面极有可能上升,这将会对北极苔原带和其它无树生态系统的分布施加进一步的限制.很显然,全球现在的气候状况对北极生态系统带来的负面影响超出更新世的任何时期,很有可能是巨大的,尤其是当各种环境变化(例如紫外线B的增加,大气中氮化合物的沉积,重金属和酸污染,放射性污染物,生物栖息地破碎化)共同作用于北极生态系统时的影响也是前所未有之际.     

环境变化背景下北极生物的多样性、分布及其适应性

生物个体是对气候变化和紫外线B(UV－B)辐射变化产生反应的基础,而且这种反应会在各种时间尺度上发生.北极地区的动物、植物以及微生物种类的多样性从表面上看是低的,而且从北方针叶林到极地荒漠逐渐减少,但其原始物种却很丰富.与这种物种多样性随纬向梯度减少的趋势相反,一些空间分布范围很广的单一优势物种的优势度则呈增长趋势.全球气候变暖可能会减少该地区的物种多样性,并限制到这些物种的分布范围,尤其是在该地区生物分布的北部边缘,一些极地特有的动物和植物种类会面临着灭绝的危险.最有可能侵入苔原地带的物种是那些目前生存在极地外缘的北方地区生物.许多植物都具有自身的特征使它们能够在以下环境中生存短暂的无冰雪覆盖的生长季节,低的太阳高度角,永久冻结地带及低的土壤温度,贫乏的养分获取条件以及极少的物理扰动.以上这些特征有些可能会限制当地物种对气候变暖的反应,但其最主要的因素是这些物种与那些潜在的入侵物种相比缺乏竞争能力.北极地区陆生动物拥有许多适应特性,这使它们能够适应北极地区剧烈的温度变化.许多动物通过冬眠或迁移来逃避极地地区的恶劣天气和资源短缺.北极地区动物生存的生物环境则相对简单几乎没有天敌、竞争者、疾病、寄生生物,但同时食物资源也很短缺.极地陆生动物可能对由气候变化带来的温暖而干旱的夏季非常不适应,这种变化将会影响到动物的迁移路线、途中栖息地,并会改变冬季积雪的状况和冻融的循环过程.气候变化还会改变动物繁殖和发育的季节,并会引来新的竞争者、捕食者、寄生生物以及疾病等.极地微生物也能很好地适应该地区的气候一些微生物甚至在-39℃的低温下还能进行代谢活动.蓝藻细菌和藻类生物有着很广泛的适应策略,这能够使它们避免(至少可以减少)紫外线的伤害.微生物能够忍受许多环境条件,而且其生长周期很短,这些特点将使它们能很快适应新的生存环境.与此形成对比的是,极地植物和动物很可能通过改变其分布范围而不是积极的生物进化来适应环境的变暖.     

不确定性及建议

在北极气候影响评估(ACIA)中,一项气候及紫外线B辐射变化对北极陆地生态系统影响的评估着重强调了预期中的变暖的深远意义,尤其是变暖在未来生态系统功能、生物多样性,以及对气候的反馈方面的深远意义.然而,尽管在有些地理区域和有些学科,我们目前对气候和紫外线B辐射驱动下生态过程及生态变化的了解已算充分,但在其他区域及学科,这种了解是微弱的.即使随着北极地区研究成果的积累和新技术的引用,近来我们的预测能力已空前提高,我们目前的了解程度也还受到各种各样的不确定性的限制.这项评估是在每一种都含有不确定性的一系列方式方法,以及常常远非完整的数据集的基础上完成的.不确定性从各种方法和概念框架而来,从不可预测的意外事件、从对模型的缺乏验证、从未来温室气体排放及气候变化的一些特别情景的应用而不是预测而来.此项评估中,以减少不确定性为目的的建议比比皆是,而且涉及所有学科.然而,一再出现的主题是在北极这样人烟稀少的偏远地区,环境变化及其影响的实验、观察及监测活动达到足够的空间广度和时间长度是极其重要的.     

在物种水平上生物对气候和紫外线B辐射预期变化的响应

控制实验表明,不同物种对每个环境因子变化变量产生的响应也存在着差异.植物往往对营养元素的变化反应最为强烈,尤其是氮素的增加.夏季增温实验表明,木本植物对温度的升高表现出了积极的响应,而地衣、苔藓类植物的丰富度却因增温而降低.物种对增温的响应主要受水分有效性和雪覆盖程度控制.在气候保持湿润的情况下,伴随着夏季温度的升高,许多无脊椎动物种群的数量都有所增加.实验表明,CO2浓度和紫外线B(UV-B)辐射的增加对植物和动物影响较小,但是,一些微生物和真菌却对紫外线B辐射的增加非常敏感,甚至可能会因此产生一些诱导突变而引起流行传染病的爆发.苔原土壤的加温、CO2浓度的升高以及矿物质营养的改善一般都会增加微生物的活动.在温带气候中,藻类往往比蓝藻细菌更占优势.冬季结冰-解冻过程的增加会导致冻壳的形成,从而会大大降低许多陆生动物的冬季存活率,改变这些动物群体的动态过程.厚的积雪会使驯鹿等植食性动物很难采食到雪下的草类植物,同时也不利于其逃避食肉动物的追捕.而无雪期的提前到来则可能会加速植物的生长.物种对气候变化的响应最初可能出现在亚种这一水平上一个具有很高遗传/群系多样性的北极植物或动物物种,演化历史已经使其具有一种适应不同环境条件的能力,这将使它们能够很快适应未来的环境变化.本土知识(IK)、航空照片和卫星图像表明一些物种的分布已经发生了变化北极植被更加趋向灌木化,而且生长也更加旺盛；北极驯鹿的分布范围最近也发生了变化；一些原来在树线以南区域活动的害虫和鸟类也在北极被发现.与此相对应,大多数在北极地区进行繁殖鸟类的数量却都在下降.根据一些模型的预测,随着气候的变暖,苔原带鸟类的数量将会大幅度地下降.据物种-气候响应模型预测,由于受到气候变暖的影响,北极地区现有物种在未来的潜在分布范围都将大大缩小和向北退缩,而一些无脊椎动物和微生物则很可能会迅速向北扩展到北极地区.     

全球变化和北方森林:阈值,移动态或逐渐变化?

21世纪所表现出来的北方气候变化的巨大性忌是引起植被的变化,其变化之大足以引起重要的社会影响.但植被变化的速度和格局却难以预测.我们评述了一些能够表明在北方森林限界上或那些种子散播限制了物种分布的地区,植被是逐渐变化的证据.但是,在一个物种分布区的中心,森林组成对气候变化是相当有弹性的,除非超过了某些阈值.在阈值点上,变化是迅速且预想不到的,常常转换至截然不同的生态系统类型.很多这类变化的诱因易受管理的影响,这说明我们在今后几十年的政策抉择将对目前气候变化的生态社会后果产生重大影响.     

Sinks for nitrogen inputs in terrestrial ecosystems: a meta-analysis of 15N tracer field studies

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.     

Net Carbon Exchange Across the Arctic Tundra-Boreal Forest Transition in Alaska 1981–2000

Shifts in the carbon balance of high-latitude ecosystems could result from differential responses of vegetation and soil processes to changing moisture and temperature regimes and to a lengthening of the growing season. Although shrub expansion and northward movement of treeline should increase carbon inputs, the effects of these vegetation changes on net carbon exchange have not been evaluated. We selected low shrub, tall shrub, and forest tundra sites near treeline in northwestern Alaska, representing the major structural transitions expected in response to warming. In these sites, we measured aboveground net primary production (ANPP) and vegetation and soil carbon and nitrogen pools, and used these data to parameterize the Terrestrial Ecosystem Model. We simulated the response of carbon balance components to air temperature and precipitation trends during 1981–2000. In areas experiencing warmer and dryer conditions, Net Primary Production (NPP) decreased and heterotrophic respiration (R _H ) increased, leading to a decrease in Net Ecosystem Production (NEP). In warmer and wetter conditions NPP increased, but the response was exceeded by an increase in R _H ; therefore, NEP also decreased. Lastly, in colder and wetter regions, the increase in NPP exceeded a small decline in R _H , leading to an increase in NEP. The net effect for the region was a slight gain in ecosystem carbon storage over the 20 year period. This research highlights the potential importance of spatial variability in ecosystem responses to climate change in assessing the response of carbon storage in northern Alaska over the last two decades.     

北极4个亚区的影响分析

在评价北极陆地生态系统影响时,人们常常强调物种和生态系统对环境变化响应的地理变化,这种变化往往与气候、生物多样性、植被带、生态系统结构和功能的南-北梯度相关联,可是,环境、生态系统的功能和结构上,以及环境史和当前气候变化的明显东-西变化显然也很重要.尽管一些地方变得温暖,但另一些地方却降温了,海洋、群岛和山脉等地理屏障的东西差异过去也对物种和植被带响应气候变化而改变分布区的能力产生了很大影响,同时,这些地理屏障为种群遗传分化和生物多样性热点区的形成提供了必要的隔离条件,这些屏障在未来气候变暖时,也将影响物种重新分布的能力.为了说明这种东西向的变化,同时也避免过分笼统或过于专业化,基于大尺度的天气和气候形成因素,北极气候影响评价项目确定了4个主要亚区.通过模拟与4个北极气候影响评价亚区有关的主要信息,导致物种分布区发生改变的地理屏障,特别是大陆的分布和海洋产生的隔离,明显会影响植被带的向北移动.对植被区向北移动的地理限制或者促进将影响将来碳的贮存和释放,以及生物圈与大气之间水和能量的交换.此外,气候变化使受威胁物种数量在各个亚区之间差别很大(白令海地区别尤其是热点),各个植被亚区重新分布的能力差异将影响每个区的生物多样性.总而言之,亚区分析表明,在整个北极地区水平上概括生态系统结构和功能的反应、物种的丧失,以及生物圈对气候系统的反馈的趋势是困难的,说明需要对北极陆地生态系统对于气候变化响应的空间变化性有深刻的认识.