Explaining the excess of rare species in natural species abundance distributions

The observation that a few species in ecological communities are exceptionally abundant, whereas most are rare, prompted the development of species abundance models. Nevertheless, despite the large literature on the commonness and rarity of species inspired by these pioneering studies, some widespread empirical patterns of species abundance resist easy explanation. Notable among these is the observation that in large assemblages there are more rare species than the log normal model predicts. Here we use a long-term (21-year) data set, from an estuarine fish community, to show how an ecological community can be separated into two components. Core species, which are persistent, abundant and biologically associated with estuarine habitats, are log normally distributed. Occasional species occur infrequently in the record, are typically low in abundance and have different habitat requirements; they follow a log series distribution. These distributions are overlaid, producing the negative skew that characterizes real data sets.     

Stability of forest biodiversity

Two hypotheses to explain potentially high forest biodiversity have different implications for the number and kinds of species that can coexist and the potential loss of biodiversity in the absence of speciation. The first hypothesis involves stabilizing mechanisms, which include tradeoffs between species in terms of their capacities to disperse to sites where competition is weak, to exploit abundant resources effectively and to compete for scarce resources. Stabilization results because competitors thrive at different times and places. An alternative, 'neutral model' suggests that stabilizing mechanisms may be superfluous. This explanation emphasizes 'equalizing' mechanisms, because competitive exclusion of similar species is slow. Lack of ecologically relevant differences means that abundances experience random 'neutral drift', with slow extinction. The relative importance of these two mechanisms is unknown, because assumptions and predictions involve broad temporal and spatial scales. Here we demonstrate that predictions of neutral drift are testable using palaeodata. The results demonstrate strong stabilizing forces. By contrast with the neutral prediction of increasing variance among sites over time, we show that variances in post-Glacial tree abundances among sites stabilize rapidly, and abundances remain coherent over broad geographical scales.     

Variance in ecological consumer-resource interactions

Food-web models use the effect size of trophic interactions to predict consumer-resource dynamics. These models anticipate that strong effects of consumers increase spatial and temporal variability in abundance of species, whereas weak effects dampen fluctuations. Empirical evidence indicates that opposite patterns may occur in natural assemblages. Here I show that spatial variance in the distribution of resource populations is sensitive to changes in the variance of the trophic interaction, in addition to the mean effect of consumers, relative to other causes of spatial variability. Simulations indicate that both strong and weak direct effects of consumers can promote spatial variability in abundance of resources, but only trophic interactions with a large mean effect size can reduce variation. Predictions of the model agree with the results of repeated field experiments and are consistent with data from published consumer-resource interactions, proving to be robust across widely varying environmental conditions and species' life histories. Thus, food-web models that embody variance in trophic interactions may have increased capacity to explain the wide range of effects of consumers documented in empirical studies.     

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.     

聚果榕小蜂群落在中国西双版纳和缅甸曼德勒之间的地理变异

榕树及其传粉榕小蜂之间的相互作用是互惠共生的经典例子，每种雌雄同株的榕树还庇护着多样的非传粉小蜂，并共生形成多营养级的榕小蜂群落，是研究群落生态学的有趣模型。区域的物种库、物种的生物地理特征及种间相互作用可能影响榕小蜂群落，但研究很少。本研究选择分布于中国西双版纳和缅甸曼德勒的雌雄同株聚果榕小蜂群落作为研究对象，比较地理分布差异对榕小蜂个体大小及共存网络的影响。结果显示：聚果榕小蜂群落在两地由相同的6个种组成，但分布于西双版纳的6种榕小蜂的个体均显著大于分布在曼德勒的同种榕小蜂。在两个分布地传粉榕小蜂均为优势种，然而在西双版纳传粉榕小蜂种群较大，而在曼德勒非传粉小蜂更丰富。在两地聚果榕有不同的榕小蜂群落结构和物种丰富度，且榕小蜂共存的网络也存在变异，在两地共存网络有相同的节点、边、连接度和全局聚类系数，但是在西双版纳的共存网络有较大的直径和较高的群落矩阵温度，这表明分布于西双版纳的聚果榕小蜂群落种间互作不太紧密，群落受到的干扰较大。由此，阐明了由地理分布所驱动的榕小蜂群落变异。     

不同放牧梯度上草原群落空间异质性的比较研究

以内蒙古呼伦贝尔草原为研究对象,利用幂函数法则定量地分析在轻度放牧、中度放牧和重度放牧3种不同利用方式下,群落的物种组成、物种多样性、群落空间异质性等群落特征。结果表明:随着放牧强度的增加,群落组成结构改变,群落丰富度逐渐降低,轻牧＞中牧＞重牧;不同放牧梯度上,各物种种群在实际分布上均具有较强的空间异质性,即偏离随机分布呈现集中分布趋势;不同放牧梯度上,群落总体空间异质性指数变化规律为轻牧=重牧＞中牧,均匀度指数的变化规律为中牧＞轻牧＞重牧。     

Trophic cascades across ecosystems

Predation can be intense, creating strong direct and indirect effects throughout food webs. In addition, ecologists increasingly recognize that fluxes of organisms across ecosystem boundaries can have major consequences for community dynamics. Species with complex life histories often shift habitats during their life cycles and provide potent conduits coupling ecosystems. Thus, local interactions that affect predator abundance in one ecosystem (for example a larval habitat) may have reverberating effects in another (for example an adult habitat). Here we show that fish indirectly facilitate terrestrial plant reproduction through cascading trophic interactions across ecosystem boundaries. Fish reduce larval dragonfly abundances in ponds, leading to fewer adult dragonflies nearby. Adult dragonflies consume insect pollinators and alter their foraging behaviour. As a result, plants near ponds with fish receive more pollinator visits and are less pollen limited than plants near fish-free ponds. Our results confirm that strong species interactions can reverberate across ecosystems, and emphasize the importance of landscape-level processes in driving local species interactions.     

Interference among deleterious mutations favours sex and recombination in finite populations

Sex and recombination are widespread, but explaining these phenomena has been one of the most difficult problems in evolutionary biology. Recombination is advantageous when different individuals in a population carry different advantageous alleles. By bringing together advantageous alleles onto the same chromosome, recombination speeds up the process of adaptation and opposes the fixation of harmful mutations by means of Muller's ratchet. Nevertheless, adaptive substitutions favour sex and recombination only if the rate of adaptive mutation is high, and Muller's ratchet operates only in small or asexual populations. Here, by tracking the fate of modifier alleles that alter the frequency of sex and recombination, we show that background selection against deleterious mutant alleles provides a stochastic advantage to sex and recombination that increases with population size. The advantage arises because, with low levels of recombination, selection at other loci severely reduces the effective population size and genetic variance in fitness at a focal locus (the Hill-Robertson effect), making a population less able to respond to selection and to rid itself of deleterious mutations. Sex and recombination reveal the hidden genetic variance in fitness by combining chromosomes of intermediate fitness to create chromosomes that are relatively free of (or are loaded with) deleterious mutations. This increase in genetic variance within finite populations improves the response to selection and generates a substantial advantage to sex and recombination that is fairly insensitive to the form of epistatic interactions between deleterious alleles. The mechanism supported by our results offers a robust and broadly applicable explanation for the evolutionary advantage of recombination and can explain the spread of costly sex.     

Introduced species and their missing parasites

Damage caused by introduced species results from the high population densities and large body sizes that they attain in their new location. Escape from the effects of natural enemies is a frequent explanation given for the success of introduced species. Because some parasites can reduce host density and decrease body size, an invader that leaves parasites behind and encounters few new parasites can experience a demographic release and become a pest. To test whether introduced species are less parasitized, we have compared the parasites of exotic species in their native and introduced ranges, using 26 host species of molluscs, crustaceans, fishes, birds, mammals, amphibians and reptiles. Here we report that the number of parasite species found in native populations is twice that found in exotic populations. In addition, introduced populations are less heavily parasitized (in terms of percentage infected) than are native populations. Reduced parasitization of introduced species has several causes, including reduced probability of the introduction of parasites with exotic species (or early extinction after host establishment), absence of other required hosts in the new location, and the host-specific limitations of native parasites adapting to new hosts.     

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.     

Chaos in a long-term experiment with a plankton community

Mathematical models predict that species interactions such as competition and predation can generate chaos. However, experimental demonstrations of chaos in ecology are scarce, and have been limited to simple laboratory systems with a short duration and artificial species combinations. Here, we present the first experimental demonstration of chaos in a long-term experiment with a complex food web. Our food web was isolated from the Baltic Sea, and consisted of bacteria, several phytoplankton species, herbivorous and predatory zooplankton species, and detritivores. The food web was cultured in a laboratory mesocosm, and sampled twice a week for more than 2,300 days. Despite constant external conditions, the species abundances showed striking fluctuations over several orders of magnitude. These fluctuations displayed a variety of different periodicities, which could be attributed to different species interactions in the food web. The population dynamics were characterized by positive Lyapunov exponents of similar magnitude for each species. Predictability was limited to a time horizon of 15-30 days, only slightly longer than the local weather forecast. Hence, our results demonstrate that species interactions in food webs can generate chaos. This implies that stability is not required for the persistence of complex food webs, and that the long-term prediction of species abundances can be fundamentally impossible.     

松嫩平原两种群落虎尾草种群有性繁殖的比较

通过调查与测定,分析了松嫩平原两种群落虎尾草种群有性繁殖的特点。结果表明,虎尾草种群的有性繁殖力较强,羊草＋虎尾草群落中虎尾草的生殖株、穗和果实的生长发育均受到种间竞争的制约,籽实生产量明显低于虎尾草群落;不同群落的虎尾草种群生物量分配规律相似,均将较多能量分配给生殖生长,产生大量种子,体现了一年生植物尽可艰保留大量后代这一繁殖策略。     

广东南岭自然保护区广东松群落特征的研究

应用植被调查样地法研究了广东南岭自然保护区广东松群落的种类组成、区系地理成分、物种多样性,以及优势种种群的年龄结构等特征.结果表明,在3 000 m2样方中,有维管植物171种,隶属于64科105属.群落的数量优势科为蔷薇科、杜鹃花科、壳斗科、山茶科、樟科、冬青科、茜草科、山矾科和木兰科等,种类组成的科、属的地理成分以热带成分占优势,温带成分也占有很大的比例,特别是群落的主要建群种以温带成分为主,体现出广东松群落具有明显的中亚热带山地针-阔叶混交林植被的典型特征.群落乔木层优势种为广东松、五列木、甜椎、疏齿木荷和福建柏等.群落的Simpson和Shannon-Wiener多样性指数分别为0.927和5.225.广东松种群的立木结构表明,I级更新幼苗少,属于相对稳定至中袁种群,将来可能被阔叶树种群取代.     

广西喀斯特地区黄枝油杉林的生境及种群结构特征

为调查珍贵树种黄枝油杉Keteleeria calcarea在广西的生境状况和种群结构特征,保护并可持续利用该物种,本研究采用样地调查法对黄枝油杉群落进行调查,分析其生境状况、土壤营养成分、伴生植物组成、种群结构特征等。调查结果显示,黄枝油杉在广西主要分布于喀斯特石山地区,从山顶至山底均有分布,亦可见于石山底部与土山相连的土石交错区,常见于阳坡或半阳坡,分布区土壤为黑色石灰土、棕色石灰土或黄土,呈酸性、中性或弱碱性,有机质和全氮含量较高,有效磷含量较低。黄枝油杉群落植被状况良好,可分为乔木层、灌木层和草本层,层间植物丰富,不同群落在物种组成上存在较大差异;黄枝油杉为乔木层的优势种,一般呈集群分布,偶见少数几株或单株分布于路边或房前屋后;少数种群因设有保护小区而种群状态较好,但是仍有部分种群人为干扰较为严重。不同种群个体的高度结构、冠幅结构和径级结构存在一定差异,恭城三江和富川麦岭种群以中老龄个体较多,融安泗顶种群以中龄个体居多,而临桂二塘种群以中小个体占优势,4个种群均存在幼苗缺乏、种群更新能力弱的情况,呈衰退趋势。生境的破坏、资源的过度采伐、结实植株数量少以及林下幼苗更新困难是黄枝油杉濒危的重要原因。因此,应加强对黄枝油杉资源和生境的保护,同时加强相关科研工作及黄枝油杉的推广应用。     

紫斑百合居群核型变异式样

分析研究云南3个紫斑百合居群核型变异，应用常规统计方法和巢式分析方法，在常规染色体水平上探讨紫斑百合居群间、个体间、细胞间和同源染色体间的变异式样。用巢式方差等级分析表明，紫斑百合染色体结构变异有4．5％左右来源于居群间，有95．5%左右来源于居群内个体间、细胞间和同源染色体间。     

异速生长关系在生物学中的应用

综述了异速生长关系在动物代谢和植物生长方面的应用．分析了异速生长关系的机理解释模型——代谢生态学理论——在生物学研究中的现状和存在的问题．该理论认为,生物的代谢速率将正比于个体质量3／4次幂,然后将生物体的其他属性同代谢速率相联系,并进行一系列的时间、空间和组织水平的尺度转换,从而代谢生态学理论可以解释个体生长、发育、种群动态、分子进化,以及物种多样性等问题．     

Rapid responses of British butterflies to opposing forces of climate and habitat change.

Habitat degradation and climate change are thought to be altering the distributions and abundances of animals and plants throughout the world, but their combined impacts have not been assessed for any species assemblage. Here we evaluated changes in the distribution sizes and abundances of 46 species of butterflies that approach their northern climatic range margins in Britain-where changes in climate and habitat are opposing forces. These insects might be expected to have responded positively to climate warming over the past 30 years, yet three-quarters of them declined: negative responses to habitat loss have outweighed positive responses to climate warming. Half of the species that were mobile and habitat generalists increased their distribution sites over this period (consistent with a climate explanation), whereas the other generalists and 89% of the habitat specialists declined in distribution size (consistent with habitat limitation). Changes in population abundances closely matched changes in distributions. The dual forces of habitat modification and climate change are likely to cause specialists to decline, leaving biological communities with reduced numbers of species and dominated by mobile and widespread habitat generalists.     

Intraspecific competition favours niche width expansion in Drosophila melanogaster

Ecologists have proposed that when interspecific competition is reduced, competition within a species becomes a potent evolutionary force leading to rapid diversification. This view reflects the observation that populations invading species-poor communities frequently evolve broader niches. Niche expansion can be associated with an increase in phenotypic variance (known as character release), with the evolution of polymorphisms, or with divergence into many species using distinct resources (adaptive radiation). The relationship between intraspecific competition and diversification is known from theory, and has been used as the foundation for some models of speciation. However, there has been little empirical proof that niches evolve in response to intraspecific competition. To test this hypothesis, I introduced cadmium-intolerant Drosophila melanogaster populations to environments containing both cadmium-free and cadmium-laced resources. Here I show that populations experiencing high competition adapted to cadmium more rapidly than low competition populations. This provides experimental confirmation that competition in a population can drive niche expansion onto new resources for which competition is less severe.     

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.