Fishing elevates variability in the abundance of exploited species

The separation of the effects of environmental variability from the impacts of fishing has been elusive, but is essential for sound fisheries management. We distinguish environmental effects from fishing effects by comparing the temporal variability of exploited versus unexploited fish stocks living in the same environments. Using the unique suite of 50-year-long larval fish surveys from the California Cooperative Oceanic Fisheries Investigations we analyse fishing as a treatment effect in a long-term ecological experiment. Here we present evidence from the marine environment that exploited species exhibit higher temporal variability in abundance than unexploited species. This remains true after accounting for life-history effects, abundance, ecological traits and phylogeny. The increased variability of exploited populations is probably caused by fishery-induced truncation of the age structure, which reduces the capacity of populations to buffer environmental events. Therefore, to avoid collapse, fisheries must be managed not only to sustain the total viable biomass but also to prevent the significant truncation of age structure. The double jeopardy of fishing to potentially deplete stock sizes and, more immediately, to amplify the peaks and valleys of population variability, calls for a precautionary management approach.     

Generation time and temporal scaling of bird population dynamics

Theoretical studies have shown that variation in density regulation strongly influences population dynamics, yet our understanding of factors influencing the strength of density dependence in natural populations still is limited. Consequently, few general hypotheses have been advanced to explain the large differences between species in the magnitude of population fluctuations. One reason for this is that the detection of density regulation in population time series is complicated by time lags induced by the life history of species that make it difficult to separate the relative contributions of intrinsic and extrinsic factors to the population dynamics. Here we use population time series for 23 bird species to estimate parameters of a stochastic density-dependent age-structured model. We show that both the strength of total density dependence in the life history and the magnitude of environmental stochasticity, including transient fluctuations in age structure, increase with generation time. These results indicate that the relationships between demographic and life-history traits in birds translate into distinct population dynamical patterns that are apparent only on a scale of generations.     

Concurrent density dependence and independence in populations of arctic ground squirrels

No population increases without limit. The processes that prevent this can operate in either a density-dependent way (acting with increasing severity to increase mortality rates or decrease reproductive rates as density increases), a density-independent way, or in both ways simultaneously. However, ecologists disagree for two main reasons about the relative roles and influences that density-dependent and density-independent processes have in determining population size. First, empirical studies showing both processes operating simultaneously are rare. Second, time-series analyses of long-term census data sometimes overestimate dependence. By using a density-perturbation experiment on arctic ground squirrels, we show concurrent density-dependent and density-independent declines in weaning rates, followed by density-dependent declines in overwinter survival during hibernation. These two processes result in strong, density-dependent convergence of experimentally increased populations to those of control populations that had been at low, stable levels.     

Why fishing magnifies fluctuations in fish abundance

It is now clear that fished populations can fluctuate more than unharvested stocks. However, it is not clear why. Here we distinguish among three major competing mechanisms for this phenomenon, by using the 50-year California Cooperative Oceanic Fisheries Investigations (CalCOFI) larval fish record. First, variable fishing pressure directly increases variability in exploited populations. Second, commercial fishing can decrease the average body size and age of a stock, causing the truncated population to track environmental fluctuations directly. Third, age-truncated or juvenescent populations have increasingly unstable population dynamics because of changing demographic parameters such as intrinsic growth rates. We find no evidence for the first hypothesis, limited evidence for the second and strong evidence for the third. Therefore, in California Current fisheries, increased temporal variability in the population does not arise from variable exploitation, nor does it reflect direct environmental tracking. More fundamentally, it arises from increased instability in dynamics. This finding has implications for resource management as an empirical example of how selective harvesting can alter the basic dynamics of exploited populations, and lead to unstable booms and busts that can precede systematic declines in stock levels.     

Collapse and recovery of marine fishes

Overexploitation and subsequent collapse of marine fishes has focused attention on the ability of affected populations to recover to former abundance levels and on the degree to which their persistence is threatened by extinction. Although potential for recovery has been assessed indirectly, actual changes in population size following long-term declines have not been examined empirically. Here I show that there is very little evidence for rapid recovery from prolonged declines, in contrast to the perception that marine fishes are highly resilient to large population reductions. With the possible exception of herring and related species that mature early in life and are fished with highly selective equipment, my analysis of 90 stocks reveals that many gadids (for example, cod, haddock) and other non-clupeids (for example, flatfishes) have experienced little, if any, recovery as much as 15 years after 45-99% reductions in reproductive biomass. Although the effects of overfishing on single species may generally be reversible, the actual time required for recovery appears to be considerable. To exempt marine fishes from existing criteria used to assign extinction risk would be inconsistent with precautionary approaches to fisheries management and the conservation of marine biodiversity.     

高寒草甸植物群落中物种多样性与种群变异性关系及其机制初探

对高寒草甸植物群落中36个物种的物种多样性与种群变异性的关系的研究结果表明:随着物种多样性的增加,种群变异性显著降低的有4个,显著增加的有1个,其余30个物种则表现出不显著的物种多样性-种群变异性的关系;1/4的物种随着多样性的增加平均种群生物量都有下降的趋势,这提供了自然群落中密度补偿存在的实验证据,而且几乎所有物种的均值-方差尺度系数Z值为1～2.这两方面的证据都与理论预期的种群变异性随多样性增加而增加的趋势不完全一致.这可能表明:多样性与种群变异性的关系随物种而异,或者说对群落中多样性与种群变异性关系的规律性认识目前的理论还无法做到,需要寻求新的模型来解释.     

Species interactions can explain Taylor's power law for ecological time series

One of the few generalities in ecology, Taylor's power law, describes the species-specific relationship between the temporal or spatial variance of populations and their mean abundances. For populations experiencing constant per capita environmental variability, the regression of log variance versus log mean abundance gives a line with a slope of 2. Despite this expectation, most species have slopes of less than 2 (refs 2, 3-4), indicating that more abundant populations of a species are relatively less variable than expected on the basis of simple statistical grounds. What causes abundant populations to be less variable has received considerable attention, but an explanation for the generality of this pattern is still lacking. Here we suggest a novel explanation for the scaling of temporal variability in population abundances. Using stochastic simulation and analytical models, we demonstrate how negative interactions among species in a community can produce slopes of Taylor's power law of less than 2, like those observed in real data sets. This result provides an example in which the population dynamics of single species can be understood only in the context of interactions within an ecological community.     

保护区的数量和种群在集合种群水平上的续存

生境的破碎化一般被认为不利于自然种群的长期存活 .在自然保护区的设置上 ,人们一般倾向于建立一个或少数几个大的自然保护区而不是具有相同总面积的更多较小的保护区 .建立了一个既包含局域种群动态 ,又包含集合种群侵占率的模型 ,并在这两个层次上进行了计算机模拟 ,结果表明 :集合种群的存活时间随着保护区数目的增大先增大而后减小 ,即保护区的数目维持在中等大小时最有利于种群在集合种群水平上的存活 ;自然保护区大小的最优值与种群的 Allee效应有关 .     

The effect of aggressiveness on the population dynamics of a territorial bird

A central issue in ecology lies in identifying the importance of resources, natural enemies and behaviour in the regulation of animal populations. Much of the debate on this subject has focused on animals that show cyclic fluctuations in abundance. However, there is still disagreement about the role of extrinsic (food, parasites or predators) and intrinsic (behaviour) factors in causing cycles. Recent studies have examined the impact of natural enemies, although spatial patterns resulting from restricted dispersal or recruitment are increasingly recognized as having the potential to influence unstable population dynamics. We tested the hypothesis that population cycles in a territorial bird, red grouse Lagopus lagopus scoticus, are caused by delayed density-dependent changes in the aggressiveness and spacing behaviour of males. Here we show that increasing aggressiveness experimentally for a short period in autumn reduced recruitment and subsequent breeding density by 50%, and changed population trajectories from increasing to declining. Intrinsic processes can therefore have fundamental effects on population dynamics.     

非线性出生率对双斑块模型动力学行为的影响

考虑一类具有非线性出生率的单种群双斑块扩散模型，研究非线性出生率和扩散对生物种群的影响.探讨系统平衡点的存在性、局部稳定性和全局稳定性，运用数值模拟分析非线性出生率和扩散常数D₁、D₂对种群动力学的影响.研究结果表明，在一些适当的假设下，绝灭平衡点或唯一正平衡点可能全局渐近稳定，且非线性出生率和扩散会使得两个斑块内种群持久生存或灭绝.通过数值模拟，当出生率(a₁)增大或密度依赖系数(A)减小时，种群总密度会增加.当D₁减小或D₂增大时，种群总密度也会增加.以上结果表明，物种的出生率和扩散对系统动力学行为起重要的作用.     

白桦天然次生林种群生态特征的研究

采用样地调查的方法,对太岳山七里峪白桦林种群进行了年龄结构分析、编制了静态生命表、绘制了存活曲线,并对该种群的四个函数（生存函数、积累死亡函数、死亡密度函数、危险率函数）进行数量分析,结果表明：①七里峪白桦种群龄级分布,不同环境下种群结构稍有差异：种群A、种群D为增长型种群,种群B、种群C、种群E趋向于稳定型．②白桦种群存活曲线接近于DeeveyⅡ型．③积累死亡率单调上升,生存率呈单调下降,上升或下降幅度前期大于后期,说明白桦种群在前期死亡率高,而种群后期则相对稳定．④在未来20年、40年、60年和80年中,白桦种群将呈现老龄级株数先增后减的趋势,种群稳定性长期维持较为困难．     

Spatial synchronization of vole population dynamics by predatory birds

Northern vole populations exhibit large-scale, spatially synchronous population dynamics. Such cases of population synchrony provide excellent opportunities for distinguishing between local intrinsic and regional extrinsic mechanisms of population regulation. Analyses of large-scale survey data and theoretical modelling have indicated several plausible synchronizing mechanisms. It is difficult, however, to determine the most important one without detailed data on local demographic processes. Here we combine results from two field studies in southeastern Norway--one identifies local demographic mechanisms and landscape-level annual synchrony among 28 enclosed experimental populations and the other examines region-level multi-annual synchrony in open natural populations. Despite fences eliminating predatory mammals and vole dispersal, the growth rates of the experimental populations were synchronized and moreover, perfectly linked with vole abundance in the region. The fates of 481 radio-marked voles showed that bird predation was the synchronizing mechanism. A higher frequency of risky dispersal movements in slowly growing populations appeared to accelerate predation rate. Thus, dispersal may induce a feedback-loop between predation and population growth that enhances synchrony.     

Host-parasitoid associations in patchy environments

Studies of insect host-parasitoid interactions have contributed much to the consensus that spatial patchiness is important in the regulation of natural populations. A variety of theoretical models predict that host and parasitoid populations, although unstable in the absence of environmental heterogeneity, may persist at roughly steady overall densities in a patchy environment owing to variation in levels of parasitism from patch to patch. Observed patterns of parasitism, however, have a variety of forms (with variation in attack rates among patches depending directly or indirectly on host density, or showing variation uncorrelated with host density). There is some confusion about the dynamical consequences of these different forms. Here we first show how the dynamical effects of all these forms of environmental heterogeneity can be assessed by a common criterion. This 'CV2 greater than 1 rule' states that the overall population densities will remain roughly steady from generation to generation if the coefficient of variation squared (CV2) of the density of searching parasitoids in the vicinity of each host exceeds approximately unity. By partitioning CV2 into components, we show that both direct and inverse patterns of dependence on host density, and density-independent patterns, all contribute to population regulation in the same way. Second, we show how a maximum-likelihood method can be applied to the kind of field data that are usually available (that is, percentage parasitism versus local host density) to estimate the components of CV2. This analysis indicates that heterogeneity is large enough to stabilize dynamics in 9 of 34 published studies, and that density-independent heterogeneity is the main factor in most cases.     

克隆植物中国沙棘种群动态及其生物量调节机制

以沙棘人工林为材料,样地调查数据为依据,以子株种群密度表示子株出生与存活能力,对其种群密度、生物量、年龄结构、克隆器官生物量投入与分配的时间动态规律及关系进行了分析。结果显示:随着种群年龄增大,子株个体数量占种群个体总数量的比例由0提高到95.5%,子株种群密度呈先上升后下降再回升趋势; 克隆种群年龄结构随子株种群密度的改变,从增长型(稳定型)演变为衰退型再演变为增长型。然而,子株种群密度与克隆器官生物量投入及其分配之间呈极显著正相关。且在种群恢复过程中,生物量投入与分配的回升相对于子株种群密度回升具有前置效应。因此,种群通过克隆器官生物量投入与分配格局调节制约子株的出生或存活从而改变子株种群密度,而子株种群密度变动过程决定克隆种群密度消长、年龄结构演变规律。根据沙棘克隆种群密度消长规律、年龄结构演变规律及其与克隆器官生物量投入、分配调节的内在联系,可将中国沙棘人工种群动态分为3个阶段:克隆生长旺盛期,克隆器官生物量投入及其分配随着种群年龄增大而上升,克隆种群密度随之上升,年龄结构为增长型(稳定型); 克隆生长衰退期,克隆器官生物量投入及其分配随着种群年龄增大而下降,克隆种群密度随之下降,年龄结构从增长型(稳定型)演变为衰退型; 克隆生长恢复期,克隆器官生物量投入及其分配随着种群年龄的增大而回升,克隆种群密度随之回升,种群年龄结构从衰退经过恢复又演变为增长型。     

Habitat structure and population persistence in an experimental community

Understanding spatial population dynamics is fundamental for many questions in ecology and conservation. Many theoretical mechanisms have been proposed whereby spatial structure can promote population persistence, in particular for exploiter-victim systems (host-parasite/pathogen, predator-prey) whose interactions are inherently oscillatory and therefore prone to extinction of local populations. Experiments have confirmed that spatial structure can extend persistence, but it has rarely been possible to identify the specific mechanisms involved. Here we use a model-based approach to identify the effects of spatial population processes in experimental systems of bean plants (Phaseolus lunatus), herbivorous mites (Tetranychus urticae) and predatory mites (Phytoseiulus persimilis). On isolated plants, and in a spatially undivided experimental system of 90 plants, prey and predator populations collapsed; however, introducing habitat structure allowed long-term persistence. Using mechanistic models, we determine that spatial population structure did not contribute to persistence, and spatially explicit models are not needed. Rather, habitat structure reduced the success of predators at locating prey outbreaks, allowing between-plant asynchrony of local population cycles due to random colonization events.     

The buffer effect and large-scale population regulation in migratory birds

Buffer effects occur when sites vary in quality and fluctuations in population size are mirrored by large changes in animal numbers in poor-quality sites but only small changes in good-quality sites. Hence, the poor sites 'buffer' the good sites, a mechanism that can potentially drive population regulation if there are demographic costs of inhabiting poor sites. Here we show that for a migratory bird this process can apply on a country-wide scale with consequences for both survival and timing of arrival on the breeding grounds (an indicator of reproductive success). The Icelandic population of the black-tailed godwit, Limosa limosa islandica, wintering in Britain has increased fourfold since the 1970s (ref. 5) but rates of change within individual estuaries have varied from zero to sixfold increases. In accordance with the buffer effect, rates of increase are greater on estuaries with low initial numbers, and godwits on these sites have lower prey-intake rates, lower survival rates and arrive later in Iceland than godwits on sites with stable populations. The buffer effect can therefore be a major process influencing large-scale population regulation of migratory species.     

Symbiotic fungal endophytes control insect host-parasite interaction webs

Symbiotic microorganisms that live intimately associated with terrestrial plants affect both the quantity and quality of resources, and thus the energy supply to consumer populations at higher levels in the food chain. Empirical evidence on resource limitation of food webs points to primary productivity as a major determinant of consumer abundance and trophic structure. Prey quality plays a critical role in community regulation. Plants infected by endophytic fungi are known to be chemically protected against herbivore consumption. However, the influence of this microbe-plant association on multi-trophic interactions remains largely unexplored. Here we present the effects of fungal endophytes on insect food webs that reflect limited energy transfer to consumers as a result of low plant quality, rather than low productivity. Herbivore-parasite webs on endophyte-free grasses show enhanced insect abundance at alternate trophic levels, higher rates of parasitism, and increased dominance by a few trophic links. These results mirror predicted effects of increased productivity on food-web dynamics. Thus 'hidden' microbial symbionts can have community-wide impacts on the pattern and strength of resource-consumer interactions.     

鲎保育的三赢策略

鲎有活化石之称,不仅是古生物学的重要研究对象,亦是医药试剂的原料,保育鲎不仅维系了人类生存,更是促进人类文明的跃升。但近年来,鲎被人类大量捕食、采血及破坏栖息地,种群大规模下降,迫切需要得到保育。根据世界自然保护联盟(IUCN)指南,鲎的物种保育工作以划设鲎保护区、增加鲎的种群量以及促进大众参与保育为重点。研究结果显示:在中国南部沿海,中国鲎(Tachypleus tridentatus)与圆尾鲎(Carcinoscorpius rotundicauda)的稚鲎共同栖息在有红树林分布的河口,这两种鲎的食物中大约60%的碳源来自海草衍生的食物;在户外养殖条件下,稚鲎与方斑东凤螺(Babylonia areolata)混养能够加速稚鲎脱壳率并提高存活率,因而可以获得大量适合放流的稚鲎(3龄)。本文在现有的研究成果基础上提出鲎保育的三赢策略:支持有效的海岸及河口保护区经营管理,在保护区内维系鲎的种群,对大众与小区进行鲎资源的明智利用宣教。     

Parasites and climate synchronize red grouse populations

There is circumstantial evidence that correlated climatic conditions can drive animal populations into synchronous fluctuations in abundance. However, it is unclear whether climate directly affects the survival and fecundity of individuals, or indirectly, by influencing food and natural enemies. Here we propose that climate affects trophic interactions and could be an important mechanism for synchronizing spatially distributed populations. We show that in specific years the size of red grouse populations in northern England either increases or decreases in synchrony. In these years, widespread and correlated climatic conditions during May and July affect populations regionally and influence the density-dependent transmission of the gastrointestinal nematode Trichostrongylus tenuis, a parasite that reduces grouse fecundity. This in turn forces grouse populations into synchrony. We conclude that specific climatic events may lead to outbreaks of infectious diseases or pests that may cause dramatic, synchronized changes in the abundance of their hosts.     

Rapid worldwide depletion of predatory fish communities

Serious concerns have been raised about the ecological effects of industrialized fishing, spurring a United Nations resolution on restoring fisheries and marine ecosystems to healthy levels. However, a prerequisite for restoration is a general understanding of the composition and abundance of unexploited fish communities, relative to contemporary ones. We constructed trajectories of community biomass and composition of large predatory fishes in four continental shelf and nine oceanic systems, using all available data from the beginning of exploitation. Industrialized fisheries typically reduced community biomass by 80% within 15 years of exploitation. Compensatory increases in fast-growing species were observed, but often reversed within a decade. Using a meta-analytic approach, we estimate that large predatory fish biomass today is only about 10% of pre-industrial levels. We conclude that declines of large predators in coastal regions have extended throughout the global ocean, with potentially serious consequences for ecosystems. Our analysis suggests that management based on recent data alone may be misleading, and provides minimum estimates for unexploited communities, which could serve as the 'missing baseline' needed for future restoration efforts.