黄土高原马栏林区辽东栎林种内、种间竞争研究

利用Hegyi的单木竞争指数模型对黄土高原马栏林区辽东栎(Quercus liaotungensis)的种内和种间竞争强度进行了定量分析.结果显示:(1)辽东栎种内竞争强度与种间竞争强度的总和大致相等,其种内及种间竞争强度均随对象木胸径的增大而逐渐减小,其关系服从幂函数规律,竞争主要发生在胸径小于15 cm的幼树阶段.(2)各组成树种对辽东栎影响程度(竞争指数)的大小顺序为:辽东栎>油松(Pinus tabulaeformis)>白桦(Betulaplatyphylla)>山杨(Populus davidiana)>杜梨(Pyrus betulaefolia)>湖北山楂(Crataegus hupehensis)>茶条槭(Acer ginnala)>陕西鹅耳枥(Carpinus shensiensis)>葛萝槭(A.grosseri).(3)油松是该地区的次优势种,对辽东栎有较大的竞争压力,在群落发展中可能会形成以油松占优势的混交林.(4)辽东栎胸径小于15 cm时,种内种间竞争是导致大量个体死亡的主要原因;胸径超过15 cm时,人为砍伐可能是个体死亡的主要原因.研究结果认为马栏林区混交林油松的密度不宜过大,尤其是在与未成熟辽东栎种群混交时密度应合理,应对该林区的混交林进行人工抚育,及时择伐密度过大的油松,而在辽东栎胸径达到15 cm之前择伐长势较差的植株,以促使森林群落尽快趋于稳定.     

Can environmental fluctuation prevent competitive exclusion in sympatric flycatchers?

Ecology has been characterized by a central controversy for decades: namely, whether the distribution and abundance of organisms are determined by species interactions, such as competitive exclusion, or by environmental conditions. In part, this is because competitive exclusion has not been convincingly demonstrated in open, natural systems. In addition, traditional theoretical models cannot predict the outcome of competitive interactions in the presence of environmental variability. In this paper we document the limiting influence of strong interspecific competition on population dynamics and nestling mortality in a mixed population of pied flycatchers (Ficedula hypoleuca) and collared flycatchers (F. albicollis in a narrow zone of sympatry. Whereas the former species was limited mainly by interspecific competition, the latter species was limited by the concerted influences of intraspecific competition and climate. The analysis suggests a march towards competitive exclusion of the pied flycatcher during warm periods. However, competitive exclusion is apparently prohibited on a local scale because intraspecific competition among individual collared flycatchers intensifies when they are forced to cope with severe environmental conditions, promoting the temporary and local presence of pied flycatchers.     

种间竞争的一个新的数学模型—对经典的Lotka—Vol—terra竞争方程的扩充

本文根据营养动力学理论,建立了一类种间竞争的新的数学模型:它是单种群增长的Cui-Lawson模型,在种间竞争上的推广。新的种间竞争模型克服了经典的种间竞争的Lotka-Volteira方程的局限与不足,具有更广泛和复杂的行为,并在特殊条件下以Lotka-Volterra竞争方程为其特例。因此,新的种间竞争的数学模型是更一般的解释性模型,是对经典的Lotka-Voterra竞争方程的扩充。     

Stochastic and deterministic processes jointly structure tropical arthropod communities

The question of whether ecological assemblages are structured by stochastic and deterministic (e.g. interspecific competition) processes is controversial, but it is difficult to design sampling regimes and experiments that can dissect the relative importance of stochastic and deterministic processes in natural assemblages. Using null models, we tested communities of arthropod decomposers in tropical epiphytes for patterns of species co-occurrence, while controlling for habitat gradients, seasonal variations and ecological succession. When environmental conditions were controlled, our analysis showed that the communities were structured stochastically. However, analysing mixed sets of communities that were deliberately created either from two distinct heights or two successional stages revealed that communities were structured deterministically. These results confirm that habitat gradients and dispersal/competition trade-offs are capable of generating non-random patterns within decomposer arthropod communities, but reveal that when such effects are accounted for, species co-occurrence is fundamentally random.     

Interspecific competition in metapopulations

The assumptions and predictions of metapopulation models for competing species are discussed in relation to empirical studies of colonization and extinction in metapopulations. In three species of Daphnia in rockpools, interspecific competition increased local extinction rates, while no effects on colonization rates were detected. Distributional patterns were consistent with several predictions of the competition model; for example, the number of species on an island increased with the number of pools and the proportion of pools occupied by each species decreased with increasing species number. It is concluded that interspecific competition is important for the distributional dynamics of Daphnia species in rockpools, but the question whether the coexistence of these species depends on metapopulation dynamics is still unresolved. Other studies on the effects of interspecific competition on colonization and extinction rates are discussed.     

The present status of the competitive exclusion principle

Since Darwin accepted the Malthusian population theory to solve the demographic problems he thought to be logically connected with the universal operation of natural selection, the numerical processes in both populations and communities were generally supposed to be governed by competition. For interspecific relations this found expression in the 'competitive exclusion principle'. After it was shown that coexistence rather than exclusion of closely related species is the rule, this principle gradually changed into the 'competitive niche shift principle'. Recently the universality of competition has been increasingly questioned, so that other interspecific relationships (especially predation) are revaluated as possibly governing many natural population and inter-population processes.     

Phylogenesis / biotope interactions among formicidae

The development of interspecific recognition and the expression of behavioral plasticity as a function of phyletic distance and the natural environment in which the ants live were investigated.An experimental model involving artificial mixed societies was used in this study. These mixed societies were produced by combining ants from two out of the 11 investigated species. The possibilities thus ranged from situations in which phylogenetically similar ants that live in the same biotype were combined, to those in which the species differ significantly in terms both of phyletic distance and of biotope.The interspecific groups were established in a neutral environment with adult ants less than 24 hours old. The results concerned agonistic and non-agonistic behaviors during the first 15 days. Both the frequency and types of interaction were shown to vary according to the type of group. Two species that share a common habitat are less aggressive and display more non-agonistic behavior than when the species come from different habitats. On the other hand, although aggressive behaviors increase with greater phyletic distance between the species, non-agonistic interactions remain constant.As a general rule, interactions between species from differing environments seem to be determined by the phyletic distance, whereas those between species from the same biotope are controlled by interspecific competition. There is a gradual change in these interactions dependent on the duration of the mixed association.     

Developmental Time and Thorax Length Differences Between the Cactophilic Species Drosophila Buzzatii and D. Koepferae Reared in Different Natural Hosts

Drosophila buzzatii and Drosophila koepferae are two cactophilic sibling species whose ranges partially overlap in Northwestern and Western Argentina. Both species can utilize the decaying tissues of both Opuntia and columnar cacti as breeding sites. Though D. buzzatii and D. koepferae are not differentially attracted to Opuntia and columnar hosts, the composition of the communities of flies emerging from natural substrates of both cacti differed significantly in a natural population. The objective of this paper is to analyze whether intra and/or interspecific competition affects development time and thorax length in D. buzzatii and D. koepferae when both species are reared in single and mixed species culture and fed with semi-natural media prepared with fermenting materials of Opuntia sulphurea(tuna) and Trichocereus terschekii(cardón). Our results showed that both traits differ significantly between flies raised in different hosts and that differences between D. koepferae and D. buzzatii species for both thorax length and development time depend on the type of culture (mixed vs. single species). In addition, the host by type of culture interaction was significant. We also observed thorax length differences between Drosophila species and type of culture. Our present data suggest that the effect of intra and interspecific competition varied between the two traits investigated and between species. However, competition alone cannot explain the differential pattern of resource utilization shown by D. buzzatii and D. koepferae in the natural population studied.     

食物、捕食和种间竞争对东方田鼠种群动态的作用

采用2×2×2析因实验设计,在野外围栏条件下,测定食物、捕食和竞争物种黑线姬鼠(Apodemus agrarius)对东方田鼠(Microtus fortis)种群动态作用的格局.食物可利用性、捕食及种间竞争的独立作用对种群最小存活数均具有极显著的效应,除捕食与种间竞争的交互作用接近显著水平外,食物与种间竞争、食物与捕食者以及三者间的交互作用均不显著;三类外部因子对种群补充量的独立作用效应均达到极显著水平,且对种群补充量的作用具有累加效应;食物可利用性、捕食及种间竞争对种群繁殖成体的比例具有极显著的作用;三类外部因子对种群幼体与成体的比例具有极显著的作用.对种群年龄结构而言,与捕食者及种间竞争比较,食物可利用性是相对较弱的影响因子,在任何捕食与种间竞争交互作用条件下,食物的作用均不显著;三类外部因子均能显著地影响东方田鼠的体重增长率,但三者的交互作用对其影响不显著;MANOVA结果表明,捕食对成体存活率的作用最强烈,其次,为食物可利用性,种间竞争的作用最弱,但三者的交互作用效应不显著.对幼体的存活时间,除捕食的作用接近显著水平外,食物可利用性及种间竞争的作用均不显著.结果提供了食物可利用性、捕食和种间竞争对东方田鼠种群动态作用的充分证据,验证了食物、捕食和种间竞争对田鼠类种群动态具有独立或累加效应的总假设.     

The interspecific competition presents greater nutrient facilitation compared with intraspecific competition through AM fungi interacting with litter for two host plants in karst soil

AM 真菌和枯落物互作下两种喀斯特植物种间竞争较种内竞争更能促进植物养分利用 枯落物是植物养分获取和土壤养分转化的关键载体。丛枝菌根(Arbuscular mycorrhizae, AM)对植物养分摄取的影响已被广泛认知。然而，在养分亏缺的喀斯特生境中，不同竞争方式的植物如何通过AM真菌和枯落物利用养分尚不清楚。本研究对两种喀斯特适生植物构树(Broussonetia papyrifera)和云贵鹅耳枥(Carpinus pubescens)进行种内竞争和种  间竞争种植处理，并通过幼套球 囊霉(Glomus etunicatum)接种或不接种处理，以及土壤中添加或不添加两物种叶片混合枯落物处理，测定了植物生物量以及氮、磷、钾浓度等指标，研究植物的生长和养分利用。研究结果表明，AM真菌对两种植物养分摄取影响不同，AM真菌显著提高了种内和种间竞争下构树的养分摄取量，但降低了云贵鹅耳枥的养分摄取量。种间竞争下接种AM真菌，枯落物添加促进了云贵鹅耳枥对氮的摄取，抑制了构树对氮的摄取。接种AM真菌和添加枯落物条件下，种间竞争的构树对氮、磷和钾的摄取量及云贵鹅耳枥对氮的摄取量均高于种内竞争；种间竞争下两物种养分竞争力呈现明显差异，即构树对磷和钾养分竞争力显著提高，对氮则不显著；云贵鹅耳枥仅对钾的养分竞争力显著降低，对氮和磷则无显著影响。这些结果说明，在AM真菌与枯落物相互作用下，两种喀斯特植物种间竞争较种内竞争更能促进植物养分利用。     

Using coloured roots to study root interaction and competition in intercropped legumes and non-legumes

Aims Root interactions between neighbour plants represent a fundamental aspect of the competitive dynamics in pure stand and mixed cropping systems. The comprehension of such phenomena places big methodological challenges, and still needs clarification. The objectives of this work were (i) to test if a species with coloured roots can be used to examine the interaction in a legume-non-legume intercropping system; (ii) to verify the importance of initial root growth on the successive root development of mixture component plants; (iii) to test if the root interaction in the shallow layers has consequences for deep root growth and (iv) to compare the effect of intraspecific and interspecific competition on root development and biomass growth.Methods A detailed study on root growth and interaction was carried out using rhizotron tubes where two legume species were grown in pure stands or were intercropped with red beet, a variety of Beta vulgaris L. with clear red roots. Within the rhizotrons, the three species were grown either without competitors, with two plants of the same species to measure intraspecific competition or with one legume and one red beet plant to study interspecific competition. The use of mixtures where one component has clearly coloured roots, together with several scalar measurements of root depth and proliferation, allowed the measurement of the root system of each species when grown in the mixtures.Important findings The use of rhizotron tubes coupled with species with coloured roots represented a valuable method to study the belowground interaction in mixed cropping systems. The initial root growth was a very important feature for the subsequent dominance of a species and it was not related to seed dimension. Initial root growth was also important because the root interactions in the shallower soil layers were found to influence the root growth in deeper soil. The root system of the red beet showed much faster and deeper growth than that of the legumes, and made red beet the dominant component in the mixtures while the legume root system was confined to the shallower soil layer. Intraspecific competition was well tolerated by the legumes, but it was limiting for the highly competitive red beet. The outcome of root interaction between neighbour plants was confirmed to be species-specific as it changed according to the intensity of the competitive effect/response of each species of the mixture: both legumes were slightly affected by the intraspecific and highly affected by interspecific competition while red beet was more affected by intraspecific competition but strongly dominant when intercropped with legumes.     

Competition and evolutionary stability of plants in a spatially structured habitat

We modelled the population dynamics of two types of plants with limited dispersal living in a lattice structured habitat. Each site of the square lattice model was either occupied by an individual or vacant. Each individual reproduced to its neighbors. We derived a criterion for the invasion of a rare type into a population composed of a resident type based on a pair-approximation method, in which the dynamics of both average densities and the nearest neighbor correlations were considered. Based on this invasibility criterion, we showed that, when there is a tradeoff between birth and death rates, the evolutionarily stable type is the one that has the highest ratio of birth rate to mortality. If these types are different species, they form segregated spatial patterns in the lattice model in which intraspecific competitive interactions occur more frequently than interspecific interactions. However, stable coexistence is not possible in the lattice model contrary to results from completely mixed population models. This clearly shows that the casual conclusion, based on traditional well mixed population models, that different species can coexist if intraspecific competition is stronger than interspecific competition, does not hold for spatially structured population models.     

Ecological and genetic correlates of long-term population trends in the park grass experiment

The Park Grass Experiment (PGE) is the longest-observed set of experimental plant communities in existence. Although the gross composition of the vegetation was at equilibrium over the 60-yr period from 1920 to 1979, annual records show that individual species exhibited a range of dynamics. We tested two hypotheses to explain why some species initially increased and why subsequently some of these (the outbreak species) decreased again. The study was designed around eight phylogenetically independent contrasts (PICs), each containing related species with different dynamics. Our first hypothesis was that persistent increasers and outbreakers have higher intrinsic rates of natural increase than control species (species without trends), allowing them to spread when interspecific competition is reduced by drought. This was tested by measuring establishment and seed production of species in field experiments, with and without interspecific competition. Seed production in outbreak species responded more strongly to release from interspecific competition than it did in either of the other groups of species. Our second hypothesis was that outbreak species eventually declined because they lacked the genetic variation necessary to adapt to the novel habitats to which they had initially spread. We tested this by measuring mating systems and genetic diversity in persistent and outbreak species in the PGE. In seven out of seven PICs tested, the outbreak species was more selfing than its persistent relative. There was a significant positive correlation between outcrossing rate and gene diversity. These results support roles for both ecological and genetic traits in long-term dynamics.     

Co-occurrence patterns and morphological similarity of semiaquatic insects (Hemiptera: Gerromorpha) in streams of Eastern Amazonia

1. To understand how the semiaquatic bug communities are shaped, it was first verified whether there was a pattern of co-occurrence between species. It was subsequently tested whether the pattern found was related to environmental variables. Lastly, it was verified whether morphological divergence between species was present in the morphological and functional traits selected. 2. A pattern of species co-occurrence was found in the studied assemblage, related to both environmental variables and interspecific biological interactions. Pairs of species with negative, positive, and random co-occurrence were found. Traits related to predation competition, sexual competition, and interaction with the environment showed significant morphological divergence. 3. Therefore, more than one process defines species co-occurrence patterns in semiaquatic bug communities. It is suggested that environmental influence is related to species microhabitat preference. On the other hand, the morphological divergence found is related to competition and sharing of food resources.     

Replacement interaction diagrams in interspecific competition experiments

Replacement interaction diagrams offer a new approach in displaying results of interspecific competition experiments. They allow visual assessment of both actual and relative responses of two species in a replacement series to treatments arranged in a factorial array. Yield data are plotted on a semi-logarithmic scale along the vertical axis. Treatments are arranged on the horizontal axis. The degree of interspecific interference or competition in mixed populations is best assessed by comparing the species response in a mixture with that in the monoculture. In competition experiments involving a number of treatment variables, replacement interaction diagrams highlight trends in treatment responses and allow some comparisions of responses between plant frequences.     

On detecting and measuring competition in spatially structured plant communities

Recent models have shown that the development of spatial structure in plant mixtures may make strong competitive interactions between species hard to detect owing to spatial segregation of the competing species. Here we address the issue of measuring interspecific competition using a simulation based on a neighbourhood population model which assumes that both dispersal and competitive interactions are localized. Using known parameter combinations we use the model to test the power and efficiency of two approaches for detecting and measuring competition. The first approach is based on measuring the response of communities to the removal of neighbours. Measures of interspecific competition based on this approach are extremely biased by spatial segregation of species, although this bias may be partially overcome by altering the spatial scale at which the effects of removals are recorded. The second approach is based on multiple regression of per capita population growth rates on local densities of the interacting species. When dispersal is restricted, the regression approach provides accurate estimates of interspecific competition coefficients when the scale of the sampling unit (i.e. the quadrats within which plants are counted) is large compared to the scale at which interactions and dispersal occur. When seeds disperse globally the removal method performs best; the regression method fails because sampling units do not form closed dynamic systems. Our results highlight the importance of tailoring methods for detecting competition to the characteristics of the species in question. They also indicate that rapid nonmanipulative estimates of competition coefficients may be the best approach in communities where dispersal is restricted and competitive interactions localized, which is likely to be the case for the majority of plants.     

On the Advantages of Mixed‐Species Groups: Impalas Adjust Their Vigilance When Associated With Larger Prey Herbivores

Prey can obtain valuable benefits from associating with other species if heterospecifics help to detect predators or locate good food patches. In mixed‐species groups, how species respond to the presence of other species remains a poorly explored question although it might give crucial insights into mechanisms underlying the interspecific coexistence. We studied temporary mixed‐species groups of large herbivores in Hwange National Park (Zimbabwe) between the common impala (Aepyceros melampus), the focal species here, and bigger species including the plains zebra (Equus quagga), the greater kudu (Tragelaphus strepsiceros) or the blue wildebeest (Connochaetes taurinus). In the Hwange savanna, the focal and smaller species are exposed to a larger range of predators than the associated species. In this context, we investigated how impalas adjusted their vigilance with group size comparing impala‐only and mixed‐species groups and whether the identity of heterospecifics affected vigilance of impalas. Our study showed that the time impalas spent in vigilance significantly decreased with group size when they formed impala‐only groups, whereas it did not significantly vary with group size in mixed‐species groups. Moreover, in mixed‐species groups, impalas did not adjust their time spent in vigilance with the proportion of conspecifics and the identity of the associated species. Thus, the mechanism underlying the difference of impalas' behavioural adjustment of vigilance with group size between single‐ and mixed‐species groups seemed to be related to the presence but not to the number and the identity of heteropecifics. Finally, we discuss the concept that larger and dominant heterospecifics were likely to increase competition for food access, thereby forcing higher vigilance of impalas, outweighing any reduction from collective vigilance.     

Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra‐ and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five‐fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra‐ and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.     

Validation of null model tests using Neyman–Pearson hypothesis testing theory

A long-standing question in ecology is whether interspecific competition affects co-occurrence patterns of species. Null model tests of presence–absence data (NMTPAs) constitute an important approach to address the question, but different tests often give conflicting results when applied to the same data. Neyman–Pearson hypothesis testing theory provides a rigorous and well accepted framework for assessing the validity and optimality of statistical tests. Here, I treat NMTPAs within this framework, and measure the robustness and bias of 72 representative tests. My results indicate that, when restrictive assumptions are met, existing NMTPAs are adequate, but for general testing situations, the use of all existing NMTPAs is unjustified — the tests are nonrobust or biased. For many current applications of NMTPAs, restrictive assumptions appear unmet, so these results illustrate an area in which existing NMTPAs can be improved. In addition to highlighting useful improvements to existing NMTPAs, the results here provide a rigorous framework for developing improved methods.