Invasive trout increase the climatic sensitivity of zooplankton communities in naturally fishless lakes

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Higher temperatures enhance the effects of invasive sportfish on mountain zooplankton communities

1. Decades of introductions of exotic sportfish to mountain lakes around the world have impoverished them biologically, and this may be exacerbated by global warming. We assessed the current status of invasive salmonids and native zooplankton communities in 34 naturally fishless lakes along an elevational gradient, which served as an environmental proxy for the expected effects of climate change. 2. Our main goal was to explore how climate‐related variables influence the effects of stocked salmonids on the total biomass, species richness and taxonomic composition of zooplankton. We predicted that warmer conditions would dampen the negative predatory effects of exotic brook trout (Salvelinus fontinalis) on zooplankton communities because more temperate lakes contain a greater diversity of potentially tolerant species. 3. Instead, we discovered that the persistence of stocked brook trout in the warmer lakes significantly amplified total zooplankton biomass and species richness. In colder and deeper lakes, zooplankton were relatively unaffected by S. fontinalis, which however persisted better in alpine lakes than at lower elevations after stocking practices were halted over two decades ago. Warmer lake conditions and higher concentrations of dissolved organic carbon (DOC) were significant primary drivers of zooplankton species turnover, both favouring greater species diversity. 4. Our findings of an ecological surprise involving potential synergistic positive effects of climate warming and exotic trout on native zooplankton communities presents a conundrum for managers of certain national mountain parks. Present mandates to eradicate non‐native trout and return the mountain lakes to their naturally fishless state may conflict with efforts to conserve biodiversity under a rapidly changing climate.     

短期增温对西藏念青唐古拉山典型植物群落结构和稳定性的影响

青藏高原气候变暖幅度显著高于全球其他区域,深刻影响着该地区植物群落的结构和稳定性。选择西藏念青唐古拉山的三种典型植物群落(高寒草原、高寒草甸和流石滩)作为研究对象,采用开顶式增温箱(OTC)模拟增温,研究了短期增温对植物群落结构和稳定性的影响。结果表明:(1)增温改变了群落的优势物种,影响其结构组成,而对物种多样性无显著影响;(2)增温显著降低了高寒草甸的地上生物量(P < 0.05),增加地下生物量(P < 0.01),从而导致了群落地下地上生物量分配策略的改变;(3)增温降低群落中部分物种的生态位宽度,进而影响群落稳定性,其中高寒草甸变化最大,达到-66.8%。研究结果可为青藏高原高寒草地生态系统应对和适应未来气候变化提供一定科学依据。     

Rising temperature and marine plankton community dynamics: Is warming bad?

Global warming is a major threat to the natural environment worldwide with potential adverse impact on plankton community. This will ultimately lead to a change in the dynamics of aquatic food webs. In this study we used seasonally forced multi-species version of the classic Rosenzweig–MacArthur predator–prey model to understand the role and stochastic influence of increasing temperature on marine plankton. First, stable coexistence of four phytoplankton and three zooplankton species was created in a system and then the level of temperature changed to achieve our research goal. We found that the stable coexistence of phytoplankton and zooplankton was related to periodic shifts in species biomass, variation in inter-specific competition and niche configuration. Warming significantly reduced total plankton biomass and changed turnover time of a species, with gradual warming breaking the stable coexistence of phytoplankton and zooplankton. In addition, we found that warming make specialist species more vulnerable than generalist species. After adding noise, a significant variation was observed in plankton biomass and amplification of noise was higher for phytoplankton compared to zooplankton. These results suggest that stochastic or unpredictable nature of temperature fluctuations may create a window of opportunity for the emergence of new species. Overall, warming would induce a shift in plankton dynamics and thereby exert pressure on plankton dependent communities such as fish in the long run.     

Interactive effects of higher temperature and dissolved organic carbon on planktonic communities in fishless mountain lakes

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Interactive effects of global warming and ‘global worming’ on the initial establishment of native and exotic herbaceous plant species

The spread of exotic earthworms (‘worming’) and rising temperatures are expected to alter the biological, chemical and physical properties of many ecosystems, yet little is known about their potential interactive effects. We performed a laboratory microcosm experiment to investigate the effects of earthworms (anecic, endogeic, epigeic, or all three together) and 4°C warming on soil water content, litter turnover and seedling establishment of four native and four exotic herbaceous plant species. Warming and worming exerted independent as well as interactive effects on soil processes and plant dynamics. Warming reduced the water content of the upper soil layer, but only in the presence of earthworms. Litter removal increased in the presence of earthworms, the effect being most pronounced in the presence of anecic earthworms at ambient temperature. Exotic plant species were most influenced by earthworms (lower seedling number but higher biomass), whereas natives were most sensitive to warming (higher seedling number). This differential response resulted in significant interaction effects of earthworms and warming on abundance and richness of native relative to exotic plants as well as related shifts in plant species composition. Structural equation modeling allowed us to address possible mechanisms: direct effects of earthworms primarily affected exotic plants, whereas earthworms and warming indirectly and differentially affected native and exotic plants through changes in soil water content and surface litter. Invasive earthworms and warming are likely to interactively impact abiotic and biotic ecosystem properties. The invasion of epigeic and anecic species could select for plant species able to germinate on bare soil and tolerate drought, with the latter becoming more important in a warmer world. Thus earthworm invasion may result in simplified plant communities of increased susceptibility to the invasion of exotic plants.     

Ecological impact of introduced fish in high altitude lakes: a case of study from the European Alps

Fish introduction is a major threat to alpine lake biota leading to the loss of native species and to the degeneration of natural food-webs. This study provides an extensive investigation on the impact of the introduced fish Salvelinus fontinalis on the native communities of alpine lakes in the Gran Paradiso National Park. We compared the macroinvertebrate and zooplankton communities of six stocked and nine fishless lakes with a repeated sampling approach during the summers 2006–2009. The impact of fish presence on alpine lake fauna is often mediated by the strong seasonality governing these ecosystems, and it dramatically affects the faunal assemblage of littoral macroinvertebrates and the size, structure, and composition of the pelagic zooplankton community with a strong selective predation of the more visible taxa. Direct ecological impacts include a decrease or extinction of non-burrower macroinvertebrates and of large zooplankton species, while small zooplankton species and burrower macroinvertebrates were indirectly advantaged by fish presence. Due to the existence of a compensation between rotifers and crustaceans, fish presence does not affect total zooplankton biomass and diversity even if fish are a factor of ecological exclusion for large crustaceans. These compensatory mechanisms are a key process surrounding the impact of introduced fish in alpine lakes.     

Effect of simulated environmental change on alpine soil arthropods

The effects of environmental change on soil animal communities are poorly known. Norwegian mountains are subject to both atmospheric nitrogen deposition and increased temperature. In a nutrient poor alpine Dryas heath in south Norway, soil arthropods were studied after 4 years of simulated environmental change by warming and/or nutrient addition. Warming alone only affected three low‐density Collembola species, while nutrient addition, with or without warming, greatly changed the dominance hierarchy of the microarthropod community. Certain Collembola species with a short (1 year) life cycle and predatory Gamasina mites increased markedly in density. These groups may have been favored by increased litter production, as plant biomass and litter producing graminoids and forbs increased significantly in plots with nutrient addition and nutrient addition combined with warming. Microarthropods with a longer life cycle, such as Oribatida and certain Collembola, were generally unaffected by nutrient addition and probably need more time to respond. The number of Oribatida taxa was, however, reduced in plots with nutrient addition, both with and without warming. A ground‐living species of Coccoidea (Homoptera) declined in plots with nutrient addition and warming compared with only warming, probably due to reduced cover of its host plant Dryas. The density of Diptera larvae (Sciaridae and Chironomidae) was unaffected by the treatments. Our results show that increased nutrient availability in nutrient poor alpine soils may have large but different effects on different taxa of soil animals. Species with short life cycles reacted first. Nutrient addition and nutrient addition combined with warming resulted in several effects below ground on microarthropods as previously shown above ground on plants: Increased biomass, high dominance of a few rapid‐growing species, contrasting responses of closely related species, and a reduction in species numbers. These short‐term responses may have profound long‐term effects in this alpine ecosystem.     

Responses of Plant Community Composition and Biomass Production to Warming and Nitrogen Deposition in a Temperate Meadow Ecosystem

Climate change has profound influences on plant community composition and ecosystem functions. However, its effects on plant community composition and biomass production are not well understood. A four-year field experiment was conducted to examine the effects of warming, nitrogen (N) addition, and their interactions on plant community composition and biomass production in a temperate meadow ecosystem in northeast China. Experimental warming had no significant effect on plant species richness, evenness, and diversity, while N addition highly reduced the species richness and diversity. Warming tended to reduce the importance value of graminoid species but increased the value of forbs, while N addition had the opposite effect. Warming tended to increase the belowground biomass, but had an opposite tendency to decrease the aboveground biomass. The influences of warming on aboveground production were dependent upon precipitation. Experimental warming had little effect on aboveground biomass in the years with higher precipitation, but significantly suppressed aboveground biomass in dry years. Our results suggest that warming had indirect effects on plant production via its effect on the water availability. Nitrogen addition significantly increased above- and below-ground production, suggesting that N is one of the most important limiting factors determining plant productivity in the studied meadow steppe. Significant interactive effects of warming plus N addition on belowground biomass were also detected. Our observations revealed that environmental changes (warming and N deposition) play significant roles in regulating plant community composition and biomass production in temperate meadow steppe ecosystem in northeast China.     

喜旱莲子草在青藏高原对模拟增温的可塑性: 引入地和原产地种群的比较

气候变暖背景下植物可通过关键性状的表型可塑性来适应环境温度的增加。表型可塑性增强进化假说预测定植到新环境中的入侵植物种群具有演化出更强表型可塑性的潜力。此前对可塑性进化的研究涵盖了外来植物性状对水分条件、光照变化、土壤养分、邻体根系以及天敌防御等的响应, 而较少有研究关注增温条件下植物重要性状的可塑性进化。已有的部分研究多集中在温带和热带地区, 而较少关注入侵植物在高寒地区对增温的响应; 且研究多集中在植物生长相关性状, 较少关注功能性状和防御性状。本研究采用同质园实验比较了喜旱莲子草6个引入地(中国)种群和6个原产地(阿根廷)种群, 在西藏拉萨模拟全天增温2℃处理下的适合度性状、功能性状和防御性状的响应差异。结果表明: (1)高寒地区模拟全天增温显著提高了喜旱莲子草总生物量(+36.4%)、地上生物量(+34.5%)、贮藏根生物量(+51.4%)和毛根生物量(+33.6%), 降低了分枝强度(-19.8%)和比茎长(-30.2%); (2)模拟全天增温使引入地种群的比叶面积和黄酮含量增加, 而原产地种群则相反。这些结果表明高寒地区全天增温2℃对喜旱莲子草可能是一种有利条件。引入地种群的适合度性状对模拟全天增温2℃的响应比原产地种群更强, 而其光能利用相关性状和防御性状的响应可能提升了其在高寒地区的适合度。因此, 在未来全球气候变暖的背景下, 高寒地区温度升高可能更有利于喜旱莲子草引入地种群的定植和扩散。     

Herbivory and nutrient limitation protect warming tundra from lowland species’ invasion and diversity loss

Herbivory and nutrient limitation can increase the resistance of temperature‐limited systems to invasions under climate warming. We imported seeds of lowland species to tundra under factorial treatments of warming, fertilization, herbivore exclusion and biomass removal. We show that warming alone had little impact on lowland species, while exclusion of native herbivores and relaxation of nutrient limitation greatly benefitted them. In contrast, warming alone benefitted resident tundra species and increased species richness; however, these were canceled by negative effects of herbivore exclusion and fertilization. Dominance of lowland species was associated with low cover of tundra species and resulted in decreased species richness. Our results highlight the critical role of biotic and abiotic filters unrelated to temperature in protecting tundra under warmer climate. While scarcity of soil nutrients and native herbivores act as important agents of resistance to invasions by lowland species, they concurrently promote overall species coexistence. However, when these biotic and abiotic resistances are relaxed, invasion of lowland species can lead to decreased abundance of resident tundra species and diminished diversity.     

Experimental effect of consumer identity on the invasion success of a non-native cladoceran

An important goal of ecologists is to identify the characteristics of native communities that help to either promote or resist the invasion of non-native taxa. In this study, experimental mesocosms were used to determine how the richness and species identity of freshwater consumers influenced the invasion success of Daphnia lumholtzi (DL), a cladoceran that has successfully invaded aquatic habitats throughout the United States. Three species of native zooplankton were used to create a gradient of richness in replicated mesocosms that included all species in monoculture, all two species combinations, and all three species together. DL was then added to each mesocosm and its biomass was monitored over time. While additional research is needed to better determine how richness affects the invasion success of DL, the presence of native zooplankton resulted in reduced levels of DL biomass in all treatments relative to the zooplankton free controls. One species (Daphnia magna, DM) was able to reduce the biomass of DL to lower levels than the other two species. DL also appeared to be unable to establish or persist over time in mesocosms with DM, unlike in mesocosms with the two other species. These species–specific effects on invasion success appeared to be related to DM’s ability to reduce algal biomass to lower levels than the other two species. Therefore, while the presence of native zooplankton helps to reduce the biomass of DL once it is introduced into novel systems, the degree to which DL is able to establish and maintain populations over time within individual systems will likely depend on the presence or absence of particular species that can have particularly strong impacts on invasibility. Using such data, we may be able to predict which habitats are most likely to be invaded based on the presence or absence of individual native species.     

Warming alters the size spectrum and shifts the distribution of biomass in freshwater ecosystems

Organism size is one of the key determinants of community structure, and its relationship with abundance can describe how biomass is partitioned among the biota within an ecosystem. An outdoor freshwater mesocosm experiment was used to determine how warming of～4 °C would affect the size, biomass and taxonomic structure of planktonic communities. Warming increased the steepness of the community size spectrum by increasing the prevalence of small organisms, primarily within the phytoplankton assemblage and it also reduced the mean and maximum size of phytoplankton by approximately one order of magnitude. The observed shifts in phytoplankton size structure were reflected in changes in phytoplankton community composition, though zooplankton taxonomic composition was unaffected by warming. Furthermore, warming reduced community biomass and total phytoplankton biomass, although zooplankton biomass was unaffected. This resulted in an increase in the zooplankton to phytoplankton biomass ratio in the warmed mesocosms, which could be explained by faster turnover within the phytoplankton assemblages. Overall, warming shifted the distribution of phytoplankton size towards smaller individuals with rapid turnover and low standing biomass, resulting in a reorganization of the biomass structure of the food webs. These results indicate future environmental warming may have profound effects on the structure and functioning of aquatic communities and ecosystems.     

Five Years of Experimental Warming Increases the Biodiversity and Productivity of Phytoplankton

Phytoplankton are key components of aquatic ecosystems, fixing CO₂ from the atmosphere through photosynthesis and supporting secondary production, yet relatively little is known about how future global warming might alter their biodiversity and associated ecosystem functioning. Here, we explore how the structure, function, and biodiversity of a planktonic metacommunity was altered after five years of experimental warming. Our outdoor mesocosm experiment was open to natural dispersal from the regional species pool, allowing us to explore the effects of experimental warming in the context of metacommunity dynamics. Warming of 4°C led to a 67% increase in the species richness of the phytoplankton, more evenly-distributed abundance, and higher rates of gross primary productivity. Warming elevated productivity indirectly, by increasing the biodiversity and biomass of the local phytoplankton communities. Warming also systematically shifted the taxonomic and functional trait composition of the phytoplankton, favoring large, colonial, inedible phytoplankton taxa, suggesting stronger top-down control, mediated by zooplankton grazing played an important role. Overall, our findings suggest that temperature can modulate species coexistence, and through such mechanisms, global warming could, in some cases, increase the species richness and productivity of phytoplankton communities.     

藏北高寒草甸群落结构与物种组成对增温与施氮的响应

气候变暖和氮沉降增加作为全球环境问题,将严重影响陆地生态系统的结构与功能.研究发现,近几十年来青藏高原增温显著,其中冬季升温最明显.而已有的研究更多关注全年增温,对冬季增温研究较少.本文基于高寒草甸地区增温和氮素添加影响研究的不足,在青藏高原高寒草甸区开展模拟增温和氮添加试验,研究长期增温与氮添加对高寒草甸群落结构与物种组成的影响.试验布设于2010年7月,地点在西藏当雄高寒草甸区,共有3种增温方式:对照、全年增温、冬季增温;每种增温处理下设置5个氮素添加梯度:0、10、20、40、80 kg N·hm^-2·a^-1,系统研究气候变暖与氮添加对高寒草甸生态系统群落结构与物种组成的影响.结果表明: 2012—2014年,增温与施氮处理均显著影响群落总盖度:全年增温处理降低了群落总盖度;在不施氮处理下,冬季增温降低了群落盖度,但在施氮处理下,随着氮剂量的提高群落盖度逐渐升高.增温与施氮对不同功能群植物的影响不同,增温处理降低了禾草与莎草植物盖度,而施氮提高了禾草植物盖度.相关分析表明,植被群落总盖度与生长旺盛期土壤含水量呈正相关关系,推测在降雨较少的季节增温导致的土壤含水量降低是群落盖度降低的主要原因.半干旱区高寒草甸土壤水分主要受降雨的调控,未来气候变化情景下,降雨时空格局的改变会显著影响植被群落盖度及组成,且大气氮沉降的增加对植被群落的影响也依赖于降雨条件的变化.     

放牧和模拟增温对藏北高寒草地植物群落特征及生产力的影响

气候变化和放牧活动对草地植物物种多样性和生产力具有重要影响。为探索藏北高寒草地植物物种多样性和生产力对增温、放牧及其交互作用的响应, 于2011年在藏北高原开始建立增温实验平台, 2016年起增设放牧、增温+放牧实验, 连续2年(2016-2017年)观测了植物群落特征、群落组成、生产力和物种多样性。结果表明, 增温和放牧对高寒草地植物高度和净初级生产力具有显著交互作用。在放牧条件下, 增温对植物高度无显著影响; 但在不放牧条件下, 增温却显著增加了植物高度。在放牧条件下, 增温对净初级生产力的影响存在年际差异, 2016年增温对生产力无显著影响, 2017年增温显著降低了植物净初级生产力; 但在不放牧条件下, 增温对植物净初级生产力无显著影响。增温和放牧对高寒草地植物物种丰富度、盖度、重要值及多样性均无显著交互作用。植物盖度在增温和放牧条件下显著降低, 杂类草物种比例显著增加, 但物种多样性均无显著变化。研究表明, 增温和放牧显著改变高寒草地群落结构。未来气候变化条件下, 放牧活动加剧有可能导致高寒草地生产力降低。     

滇西北高原典型湿地湖滨带优势植物种子繁殖对增温的响应

基于IPCC对未来大气增温的预测,采用开顶式生长室(OTC)增温技术,以不增温为对照,研究滇西北高原典型湿地纳帕海湖滨带优势植物水葱和黑三棱在大气增温2.0和3.5℃处理下种子的繁殖特征.结果表明:大气增温对植物结实率影响显著,但存在种间差异.其中,增温对水葱的结实率有促进作用;黑三棱结实率在增温2.0℃处理下显著下降,但在增温3.5℃处理下又恢复到对照水平.增温促进了2种植物穗的生长,在增温2.0、3.5℃处理下水葱的穗长分别增加82.9%、89.0%,小穗数分别增加133.3%、150.0%,每株穗生物量分别增加了10.1%、89.6%,每株穗生物量占总生物量的比例分别增加79.5%、409.3%;在增温2.0、3.5℃处理下,黑三棱穗长分别增加66.1%、95.2%,每株穗生物量占总生物量的比例分别增加878.8%、1052.6%.增温显著增加了水葱和黑三棱每穗种子产量,在增温2.0、3.5℃处理下,水葱每穗种子产量分别增加33.7%、58.3%,黑三棱每穗种子产量分别显著增加3.4%、69.5%.在增温2.0、3.5℃处理下水葱种子长分别增加5.4%、6.9%,种子长宽比分别增大9.1%、5.3%;而增温对黑三棱的种子形态无显著影响.最高温和最低温是影响2种植物种子繁殖的主要温度因子.温度增高引起的生长季提前、营养生长期延长以及有机物积累量增加,为植物繁殖扩散提供了充足的物质和能量积累,可能是2种植物在增温条件下繁殖能力提高的原因.     

Responses of biotic interactions of dominant and subordinate species to decadal warming and simulated rotational grazing in Tibetan alpine meadow

Warming increases competition among plant species in alpine communities by ameliorating harsh environmental conditions, such as low temperatures. Grazing, as the main human activity, may mitigate the effect of warming, as previously reported. However, it is critical to refine the effects of warming on biotic interactions among species, for example, by taking the competitive ability of species into consideration. Based on a 10-year warming and grazing experiment in a Tibetan alpine meadow, we evaluated interspecific biotic interactions of dominant and subordinate species, using the approach of interspecific spatial associations. Warming significantly increased competition between subordinate and dominant species as well as among subordinate species, but not among dominant species. Moreover, facilitation of dominant-subordinate species also increased under warming. Simulated rotational grazing had similar effects to warming, with increasing interspecific competition. Our results show that, when studying the effects of warming on biotic interactions among species, it is necessary to characterize different species pairs relative to their competitive ability, and that simulated rotational grazing does not mitigate the effects of warming in the long term. Our results also provide evidence that the spatial pattern of species is a critical mechanism in species coexistence.     

太湖梅梁湾大型控藻围栏对浮游甲壳动物群落结构的影响

2005年对太湖梅梁湾大型鲢、鳙控藻围栏内外浮游甲壳动物群落结构的季节变化进行了监测.结果表明:围栏内外的环境因子、浮游植物生物量、浮游甲壳动物种类组成无显著差异.但鲢、鳙放养对浮游甲壳动物的生物量产生了较大的影响,围栏内浮游甲壳动物的总生物量和枝角类的生物量显著低于围栏外.总体上,枝角类各种类的生物量受鲢、鳙放养的影响程度大于桡足类的种类.太湖梅梁湾浮游甲壳动物的季节演替明显,大部分种类只是季节性出现.冬季和春季以溞(Daphnia sp.)和近邻剑水蚤(Cyclops vicnus)等大型种类为主,夏季和秋季以象鼻溞(Bosmina sp.)、角突网纹溞(Ceriodaphnia cornuta)和中华窄腹剑水蚤(Limnoithona sinensis)等小型种类为主.典范对应分析表明,透明度、温度和浮游植物的生物量是影响浮游甲壳动物季节变化的主要因素.     

模拟增温对青藏高原多年冻土区小嵩草和藏嵩草生长与抗氧化特征的影响

采用开顶式增温小室(OTCs)方法模拟气候变暖,分别选取青藏高原腹地风火山地区高寒小嵩草草甸和高寒藏嵩草沼泽草甸优势物种小嵩草和藏嵩草为研究对象,对比分析增温处理下两种优势物种叶片的形态与生理特征变化,从而探索高寒植物对气候变暖的内在响应机理.结果表明: 增温显著增加了小嵩草叶片长度(40.0%)和叶片数量(72.7%),也显著增加了藏嵩草株高(11.9%)和叶片长度(19.3%),促进了两种优势植物的形态生长和地上生物量增加.增温处理下小嵩草和藏嵩草叶片的膜透性(电导率),活性氧(过氧化氢和超氧阴离子自由基),超氧化物歧化酶、过氧化物酶、抗坏血酸过氧化物酶和过氧化氢酶活性,丙二醛含量均没有显著变化.但抗坏血酸和游离脯氨酸含量在藏嵩草叶片内分别显著增加了29.8%和53.8%,而在小嵩草叶片内没有明显变化.可见,增温下小嵩草和藏嵩草均能够维持正常的抗氧化水平,以维持该区域优势植物生长;但藏嵩草生理过程对增温更加敏感.