小麦/大豆间作中作物种间的竞争作用和促进作用

春小麦／春大豆间作是西北一熟制灌区广泛采用的高产种植形式．本文采用田间小区和微区根系分隔试验研究了这种种植形式作物种间的竞争作用和促进作用．结果表明,小麦／大豆间作具有明显的间作优势．土地当量比为１．２３～１．２６．小麦为优势种,竞争力强于大豆,具有明显的间作边行优势．小麦边行优势的１／３贡献来自于地下部．小麦收获后,大豆生长具有恢复作用,认为这种恢复作用是间作优势的机制之一．间作相对于单作两种作物的收获指数均有显著提高．收获指数的种间促进作用是间作优势的另一机制．     

Predicting extinctions: interspecific competition,predation and population variability in experimental Daphnia populations

We examine how interspecific competition and two types of size-selective predation affect population density, variability and persistence in laboratory cultures of two species of Daphnia, D. magna and D. longispina. When both species were analysed together, and for D. longispina alone, there were weak negative relationships between mean population density and population variability. Interspecific competition resulted in lower population densities and higher population variability. Extinct populations had lower densities and were also more variable than persisting ones. There was still an effect of population variability on extinction probability after the effect of density on population variability had been accounted for. Hence, the effects of population density and variability on population persistence were partly independent of each other. The effects of size-selective predation on population persistence were more species-specific and not directly related to density or variability. Since the effects of species interactions on persistence were large, we suggest that it is likely that population vulnerability analyses not incorporating effects of interspecific interactions are often misleading.     

Tree competition and species coexistence in a warm-temperate old-growth evergreen broad-leaved forest in Japan

The growth dynamics and mode of competition between adult trees 5.0cm in diameter at breast height (DBH) of nine abundant treespeciesoccupying ca. 85% of the total basal area were investigated in a 4ha study plot (200 m × 200 m) of awarm-temperate old-growth evergreen broad-leaved forest in the Tatera ForestReserve of Tsushima Island, southwestern Japan. In the plot, adult trees 5.0 cm DBH co-occurred with 35 woody plant species (except forwoody vine species). The most dominant and largest species,Castanopsis cuspidata var. sieboldiiexhibited a bimodal DBH distribution; it was found in both the upper and lowervertical layers. Other tree species had unimodal DBH distributionscorrespondingmostly to the lower vertical layer. We developed a model for individual growthincorporating both intra- and interspecific competition and degree ofcompetitive asymmetry. One-sided interspecific competition was detected in 17cases out of the 66 possible combinations on the scale of the 4 hastudy plot. The direction of interspecific competition was generally one-sidedfrom layer-I species to layer-II and III ones. The effects of two-sidedcompetition were detected only in layer-II and III species. OnlyDistylium racemosum exhibited one-sided intraspecificcompetition. We also found 11 cases of positive interspecific relationships.Generally, competitive relationships prevailed over positive relationshipsbetween adult trees in this warm-temperate evergreen broad-leaved forest.Competition between adult trees 5.0 cm in DBH did not occurinthe same vertical layer, but occurred only between trees in different verticallayers. This suggests that competition between adult trees 5.0cm in DBH plays a key role in the variation in species coexistencebetween different vertical layers on the 4 ha scale of thewarm-temperate evergreen broad-leaved forests. Moreover, it was found bycomparing with three different forest types that interspecific competition ismore intense in warm-temperate forests than in cool-temperate or sub-borealforests. We conclude that, compared to cool-temperate or sub-boreal forests(which have little interspecific competition), warm-temperate forests supportmore complex interspecific relationships and species-specific habitatpreferences that result in higher species diversity.     

Savanna tree–grass competition is modified by substrate type and herbivory

Question: Woody plant and grass interactions in savannas have frequently been studied from the perspective of the response of one growth form on the other but seldom evaluated as two‐way interactions. What causes woody plant encroachment in semi‐arid savannas and what are the competitive responses of tree seedlings and grasses on rocky and sandy substrates? Methods: In this greenhouse study, we investigated the influence of substrate and grazing on responses to interspecific competition by tree seedlings and grasses. We measured competitive/facilitative responses on biomass and nutrient status of tree seedlings and grasses grown together. Results: Interspecific competition suppressed growth of trees and grasses. Tree seedlings and uncut grass accumulated double the biomass when grown without competition relative to when they competed. Competitive responses varied on different substrates. Grass biomass on rocky substrate showed no response to tree competition, but appeared to be facilitated by trees on sandy substrate. Grass clipping resulted in higher tree seedling biomass on rocky substrate, but not on sandy substrate. There was a positive response of grass nutrient status to competition from tree seedlings. Conclusion: Selective grass herbivory in the absence of browsing or suppression of shade‐intolerant grasses by trees are commonly cited reasons behind bush encroachment in savannas. We show that grazing may confer a competitive advantage to tree seedlings and promote bush encroachment more readily on rocky substrates. This may be due to the imposed sharing of the soil depth niche on rocky substrates, whereas possible niche separation on sandy substrates minimizes the advantage conferred by reduced competition.     

Structure,organization, and response of a species-rich parasitoid community to host leafminer population dynamics

The parasitoid community dynamics of an agromyzid honeysuckle leafminer, Chromatomyia suikazurae (Agromyzidae, Diptera) were studied between 1981 and 1990 in a natural forest in Kyoto, Japan. The parasitoid fauna composed three koinobionts (all larval-pupal solitary parasitoids) and 22 idiodiont species (11 larval solitary, nine pupal solitary and one pupal gregarious). The parasitoid community was dominated by early-attacking oligophagous braconid koinobionts at early periods, but was gradually displaced by late-attacking polyphagous eulophid idiobionts. Accordingly, the diversity index of the parasitoid community peaked at an intermediate point in the intra-generational succession. The succeeding attack-in-waves by the late-attacking idiobionts greatly reduced not only the survival rates of early-attacking parasitoid larvae but also the survival rates of hosts. The density-dependence observed in the host pupal mortality was thought to result from density-dependent host-switching by a keystone polyphagous pupal idiobiont parasitoid, Chrysocharis pubens, whereas high host pupal mortality was potentially attained by an early-attacking koinobiont braconid. Supposed aggregation of polyphagous parasitoids at high host density resulted in intense within-host competition and in an increase of host-feeding attack, both of which contributed to low emergence rates of parasitoids at high host densities. Parasitoid emergence rates were also reduced at low host densities, probably by inter- and intra-specific hyperparasitism among oligophagous parasitoids for limited hosts. The regulation effects of the species-rich parasitoid community upon the host population dynamics are thought to derive from succeeding attack-in-waves by polyphagous late-attacking idiobionts, especially by the keystone species.     

Interspecific competition among macroparasites in a density-dependent host population

 We analyze the dynamics of a community of macroparasite species that share the same host. Our work extends an earlier framework for a host species that would grow exponentially in the absence of parasitism, to one where an uninfected host population is regulated by factors other than parasites. The model consists of one differential equation for each parasite species and a single density-dependent nonlinear equation for the host. We assume that each parasite species has a negative binomial distribution within the host and there is zero covariance between the species (exploitation competition). New threshold conditions on model parameters for the coexistence and competitive exclusion of parasite species are derived via invadibility and stability analysis of corresponding equilibria. The main finding is that the community of parasite species coexisting at the stable equilibrium is obtained by ranking the species according t! o th e minimum host density H ^* above which a parasite species can grow when rare: the lower H ^* , the higher the competitive ability. We also show that ranking according to the basic reproduction number Q ₀ does not in general coincide with ranking according to H ^* . The second result is that the type of interaction between host and parasites is crucial in determining the competitive success of a parasite species, because frequency-dependent transmission of free-living stages enhances the invading ability of a parasite species while density-dependent transmission makes a parasite very sensitive to other competing species. Finally, we show that density dependence in the host population entails a simplification of the portrait of possible outcomes with respect to previous studies, because all the cases resulting in the exponential growth of host and parasite populations are eliminated.. Received: 24 June 1996 / Revised version: 28 April 1998     

The relative importance of predation and competition in two reef fishes

Competition and predation may both strongly influence populations of reef fishes, but the importance of these processes relative to one another is poorly understood. I quantified the effects of predation and competition on the growth and survival of two temperate reef fishes, Lythrypnus dalli and Coryphopterus nicholsii, in field experiments in which I manipulated the densities of the two species and the abundance of predators (using exclosure cages) on small replicate patch reefs. I also evaluated the influence of predators on the behavior of the two species to help interpret the mechanisms of any predatory influences on growth or survival. Predation was much more important than competition (inter- or intraspecific) in Lythrypnus. For Coryphopterus, neither competition nor predation were particularly important. Behaviorally, both species responded to predators by reducing foraging rate and hiding. This altered behavior, however, had no repercussions for growth or survival of Coryphopterus. In contrast, Lythrypnus grew more slowly and suffered greater mortality when exposed to predators. Interspecific competition did not significantly influence either species. Intraspecific competition did not affect the growth of Coryphopterus, but survival tended to be lower at high densities. Growth of Lythrypnus was depressed by intraspecific competition, but survival was not, except that, in the presence of predators, survival was density dependent. In contrast to the historical emphasis placed on the role of competition, this study indicates that predation can be more important than competition in determining patterns of abundance of some reef fishes. For example, predators not only influenced foraging of both Lythrypnus and Coryphopterus, but they also reduced growth and survival of Lythrypnus, and therefore appear to help maintain the marked habitat segregation between the two species. Received: 16 June 1997 / Accepted: 3 December 1997     

Metapopulation structure in the lagoon cockle <Emphasis Type="Italic">Cerastoderma lamarcki</Emphasis> in the northern Wadden Sea

Benthic invertebrates in discontinuous inshore habitats and with short or no pelagic larval dispersal are likely to exhibit regional metapopulation dynamics with partially isolated local populations. Near the island of Sylt, the bivalve Cerastoderma (Cardium) lamarcki (Reeve, 1844, syn. in part with C. glaucum Bruguiére, 1789) was widespread in intertidal seagrass beds, coexisting with the sibling species C. edule (Linné, 1758). However, the last C. lamarcki in this habitat was found in 1980. At present the lagoon cockle is restricted to disjunct ditches, creeks and ponds within island salt marshes. There it differs in year-class structure between localities. Successful recruitment events did not coincide. At one locality, a period with regular recruitment was followed by 5 years of recruitment failure, resulting in an overaged population probably at the rim of extinction. In a nearby brackish pond, extinction was followed by recolonization 3 years later. Other lagoonal habitats which seem to be suitable are without cockles. It is speculated that small and isolated habitats occasionally receive colonizers by eggs and juveniles adhering to avian vectors. Electronic Publication     

Chaotic pairwise competition

We investigate a kind of competition possible in a system of at least three populations competing for the same limited resource. As a model we use generalised Volterra equations in which the growth rates and competition coefficients of populations depend on the number of members of all populations. Because of the nonconstant values of the last quantities the system could be repelled from the state of cyclic pairwise competition described by May and Leonard (SIAM J. Appl. Math. 29 (1975) 243.). We investigate the competition in a chaotic regime of evolution of the number of members of populations. We show that the nonconstant competition coefficients can lead to a regularisation of the time intervals of domination of each population and the non-constant growth rates can lead to decreasing length of the time intervals of domination as well as to chaotisation of the occurrence of these intervals. A quantity characterising the time intervals between the successive maxima of the number of the populations individuals is discussed. By means of the wavelet transform modulus maxima method we calculate the tau(q)-spectrum and the H?lder exponent for the time series of this quantity. The results of the theory are illustrated by an example of competition among the three main political parties in Bulgaria and we discuss qualitative aspects of the dynamics of change of preferences of voters.     

Pollen competition and environmental effects on hybridization dynamics between Phlox drummondii and Phlox cuspidata

Pollen competition between species strongly influences hybridization dynamics in plants. By performing single- and mixed-donor pollinations, we show that soil Ca alters the outcome of interspecific pollen competition in the annual Phlox hybrid system of Phlox cuspidata and P. drummondii. In the absence of interspecific pollen competition, heterospecific pollen siring success of both species was influenced most strongly by the maternal growth environment, such that hybridization was facilitated when heterospecific pollen was deposited on stigmas of maternal plants growing in high Ca soils. When heterospecific pollen was forced to compete against conspecific pollen, however, the maternal growth environment did not influence hybridization, but the environmental origin of heterospecific pollen did, and this effect depended on the maternal species. Pollen of P. drummondii was more effective at outcompeting P. cuspidata pollen and preventing hybridization in P. drummondii dams when P. cuspidata pollen was derived from low Ca. Pollen competition within pistils of P. cuspidata was unaffected by pollen Ca environment. In situations in which P. cuspidata grows in lower soil Ca than P. drummondii, as has been documented in one population, these results suggest that the competitive ability of heterospecific pollen will be diminished by environmental effects of soil Ca. Thus, the environment in which pollen develops can influence interspecific pollen competition and hybridization frequency.     

Relative movement patterns of a tephritid fly and its parasitoid wasps

The extent of within-patch dispersal by a tephritid fly and its four major parasitoids was examined over three field seasons. Hosts and parasitoids were marked using acrylic paint and observed as they oviposited into the flowerheads of marsh thistle, Cirsium palustre. The average recapture rate pooled across all species was 22%. The four parasitoids showed consistently greater rates of movement than the host in all three years. In nearly all comparisons, male dispersal was less than female dispersal. There was no evidence that parasitoids moved longer distances after visiting low quality rather than high quality patches. In the one season it was studied, no correlations between movement and insect size were observed. The relevance of these observations to host-parasitoid population dynamics is discussed.     

Effects of herbivory and competition on an introduced plant in decline

Anthemiscotula was introduced to Denmark 500 years ago, and its distribution is presently limited and in decline. A manipulative field experiment was performed to investigate the effects of native plant competitors and native invertebrate herbivores on its performance. Generally, both herbivory and competition treatments had great impact, and when both factors were operating, the effects were additive for all variables except plant height. Although A. cotula showed plasticity in growth, resource allocation and flowering timing, it was unable to adjust to competition and compensate for losses due to herbivory sufficiently to ensure and restore its achene production. This vulnerability, combined with improved cereal cleaning techniques and thus fewer reintroductions of A. cotula seeds, may be the cause of its current decline. A. cotula responded to herbivory by prolonging its flowering period, a “bet-hedging” strategy. In Denmark this strategy is unreliable since risks of sub-optimal conditions are much greater in August–October. Received: 12 December 1997 / Accepted: 14 October 1998     

Estimating competition coefficients: strong competition among three species of frugivorous flies

Despite the abundance of studies on competitive interactions, relatively few experiments have been used to fit explicit competition models and estimate competition coefficients. Such estimates are valuable for making contact between theoretical and empirical studies, which tend to measure competition in different units. To quantify the strength of competitive interactions among the larvae of three species of frugivorous flies, I manipulated the densities of each species to investigate all three pairwise interactions. The densities of each species were changed independently (i.e., using a response surface experimental design), which allowed maximum likelihood estimation of the competition coefficients for each species, based on the Hassell and Comins competition model. The effects of competitor density on larval survival, time to emergence, and the weight of emerging adults were also analyzed to investigate the responses of individual species to density. The estimates of the competition coefficients suggest that the larvae of these flies experience strong asymmetric competition for resources, and raise questions as to how these species coexist. For each pair, one of the species was largely unaffected by interspecific competition, but decreased the performance of the other. Received: 8 February 1999 / Accepted: 5 April 1999     

Role of the intraspecific competition in the regulation of Agrobacterium tumefaciens transconjugant population level in soil experiments

Abstract In microcosms of sterilized soil simultaneously inoculated with Pseudomonas aeruginosa carrying the plasmid R68-45 and the plasmid-free Agrobacterium tumefaciens , transconjugants were detectable after two days of incubation and their number remained constant thereafter. The growth of a transconjugant strain was monitored in sterile soil. When mixed together with the parental strains at high inoculum or when the soil was previously colonized by the donor, the transconjugant was able to grow. If the recipient was the first soil colonizer, the challenging population of transconjugant remained stable at its initial level. We demonstrated the possible role of intraspecific competition in the limitation of transconjugant numbers.     

Conditional dispersal under kin competition: extension of the Hamilton-May model to brood size-dependent dispersal

I consider a site-based model with contest competition among siblings, and assume that dispersal is conditional on the number of offspring in the natal site. Evolutionarily stable populations contain threshold dispersal strategies, which retain a certain number of offspring in the natal site and disperse the rest (if the actual number of offspring is less than the threshold, then all offspring are retained). Due to the discrete nature of the strategy set (the threshold must be integer), the ESS may not be unique or may not exist. In the latter case, two neighboring threshold strategies coexist in the evolutionarily stable population. Dispersal first decreases and then increases as a function of dispersal mortality, such that all but one offspring should be dispersed both when dispersal mortality is very small or very high. Population-level dispersal fractions are often similar to the unconditional ESS, but differ strongly when fecundity is small and dispersal mortality is high.     

Island size, isolation, or interspecific competition? The breeding distribution of the Parus guild in the Danish archipelago

The Parus guild (Parus spp., Sitta, Certhia, and Regulus) is distributed as a complex mosaic within the Danish archipelago, with from one to eight species on different islands. We assessed the roles of island isolation, island size, and interspecific competition in determining the breeding species compositions of this guild on 53 Danish islands. Small, isolated islands supported fewer species than larger, nearshore islands. These effects, however, were largely restricted to a few sedentary species (P. cristatus, P. palustris, S. europaea) that are known to be poor dispersers/colonizers. In some cases, these three species were also absent from large, nearshore islands with suitable habitat, suggesting that habitat availability was not always responsible for the absence of a species. Monte Carlo simulations suggested that the pattern of species presence/absence was not a result of interspecific interactions. Thus, although a number of previous studies have documented interspecific competition among members of the Parus guild, our results suggest that such competition is not responsible for the unusual pattern of species distribution within the Danish archipelago. Received: 28 October 1996 / Accepted: 7 February 1997     

Use of quantitative PCR to understand within-host competition between two entomopathogenic fungi

Interactions between the entomopathogenic fungi Zoophthora radicans and Pandora blunckii infecting larvae of Plutella xylostella were investigated. This is the first report to quantify within-host growth of one fungus in the presence of another competing for the same host resource using quantitative PCR (qPCR) at regular time points during the infection process. In larvae inoculated only with Z. radicans, there was a cumulative increase in the quantity of Z. radicans DNA throughout the time course of infection. However, in dual-inoculated larvae, there was an initial accelerated rate of growth of Z. radicans compared to when it was inoculated alone, but by the time of host death it had been effectively excluded by P. blunckii. The implications of these results for co-existence of these fungal pathogens in the field are discussed.     

Population biology of multispecies helminth infection: competition and coexistence

The role that interspecific interactions play in shaping parasite communities is uncertain. To date, models of competition between helminth species have assumed that interaction occurs through parasite-induced host death. To our knowledge, there has been no theoretical exploration of other forms of competition. We examine models in which competition acts at the point of establishment within the host, and at the time of egg production by the adult worm. The models used are stochastic and we allow hosts to vary in their rate of exposure to infective larvae. We derive the Lotka-Volterra model of competition when exposure is homogenous and thus demonstrate that two helminth species cannot coexist on a single limiting resource. We show that coexistence of species is promoted by heterogeneity in host exposure provided that the rates of exposure to the two species are not perfectly correlated, and, if they are positively correlated, provided that the degree of heterogeneity in host exposure is similar for the two competing helminth species. These results are robust to the mechanism of competition.     

Seasonal dynamics and interspecific competition in Oneida Lake Daphnia

I investigated the population dynamics and competitive interactions of two species of the suspension-feeding crustacean Daphnia in Oneida Lake, N.Y. Both species have persisted in the lake for decades, but their water-column densities are negatively correlated. The larger Daphnia pulicaria dominates in some years, the smaller D. galeata mendotae in others, and in some years one species replaces the other seasonally. Although this pattern results in part from annual variation in vertebrate predation pressure, predation alone cannot explain the irregular daphnid dynamics. In 1992–1995, I examined the water-column abundances, birth and death rates of both species. D. pulicaria dominated in two years, D. galeata mendotae was replaced by D. pulicaria in one year and in 1994, both species persisted in low numbers. To test the effect of temporal changes on the strength of intra- and interspecific competition on both juvenile and adult daphnids, I manipulated a series of field enclosures in 1994 and 1995. The outcome of competition varied within and between years, and its effects were most evident at the highest densities and lowest resource levels. For adults of both species, the effects of interspecific competition were detected more often than those of intraspecific competition. Lipid reserves (a metric of fitness) among juveniles were generally low, with those of D. galeata mendotae often being less than those of D. pulicaria. Contrary to the results of other studies examining competition in daphnids, spatial segregation and predictable within-year reversals in competitive dominance most likely do not play a large role in fostering coexistence of the Oneida Lake daphnids. Instead, coexistence of these competitors is promoted by interannual variation and long-lived diapausing eggs. Received: 20 July 1997 / Accepted: 21 November 1997     

Interspecific competition between insect herbivores: asymmetric competition between cinnabar moth and the ragwort seed-head fly

ABSTRACT.

• 1 Removal field experiments and observational studies have been undertaken to determine whether feeding by cinnabar moth Tyria jacobaeae L. on the flower heads of ragwort Senecio jacobaea L. affects the abundance of the fly Pegohylemyia seneciella (Meade) that feeds in the flower heads as a larva.
• 2 Correlations between the population density of cinnabar moth and the population density of the fly were suggestive of habitat separation, but provided little evidence of exploitation competition.
• 3 Removal of cinnabar moth by hand from replicated plots over two years shows that, in years when ragwort flower production is consumed by cinnabar moth caterpillars, the fly may show no recruitment at all.
• 4 Fly populations persist in refugia, exploiting ragwort plants that grow in areas where there are no cinnabar moth.
• 5 Recruitment of ragwort is not seed limited, so the reduction in seed production caused by P. seneciella (maximum about 30%) has no impact on ragwort abundance, or on the abundance of cinnabar moth.
• 6 We conclude that there is strong interspecific competition between these two species, and that the competition is highly asymmetric. The cinnabar moth had a substantial effect on the recruitment of the fly in 1986, but the fly has no measurable impact on the recruitment of the moth. In six years out of seven in our long-term study, cinnabar moth reduced flower production to levels comparable to those measured in 1986, and we infer that strong competition with the fly was likely in six years out of seven.
• 7 One reason why there are so few published examples of asymmetric interspecific competition may be simply that the experiments are thought too obvious to be worth doing. We argue that this is not a good reason for eschewing manipulative field experiments, and that few processes in ecology are at all obvious when investigated in detail.