The effect of travel time on oviposition behavior and spatial egg aggregation: experiments with Drosophila

A higher degree of spatial egg aggregation is often observed in environments where resource patches are more sparsely distributed. This suggests a higher probability of species coexistence when resource distribution is sparse. However, it is still unclear how the degree of spatial egg aggregation increases. I propose a model to explain this phenomenon, which assumes that (i) egg load (the number of mature eggs in ovaries) increases in the travel period between resource patches and (ii) the retention of eggs in the ovaries is harmful (egg load pressure). With these assumptions, a female would lay accumulated eggs on arrival at a new resource patch, resulting in a higher degree of spatial egg aggregation. Laboratory experiments with three drosophilid species, Drosophila simulans Surtevant, Drosophila auraria Peng, and Drosophila immigrans Sturtevant, support the model. This study provides evidence that host availability affects the spatial egg aggregation via egg load.     

Spatial patterns and competition of tree species in a Douglas-fir chronosequence on Vancouver Island

While the successional dynamics and large-scale structure of Douglas-fir forest in the Pacific Northwest region is well studied, the fine-scale spatial characteristics at the stand level are still poorly understood. Here we investigated the fine-scale spatial structure of forest on Vancouver Island, in order to understand how the three dominant species, Douglas-fir, western hemlock, and western redcedar, coexist and partition space along a chronosequence comprised of immature, mature, and old-growth stands. We quantified the changes in spatial distribution and association of the species along the chronosequence using the scale-dependent point pattern analyses pair-correlation function g(r) and Ripley's L-function. Evidence on intra- and inter-specific competition was also inferred from correlations between nearest-neighbor distances and tree size. Our results show that 1) the aggregation of Douglas-fir in old-growth was primarily caused by variation in local site characteristics, 2) only surviving hemlock were more regular than their pre-mortality patterns, a result consistent with strong intra-specific competition, 3) inter-specific competition declined rapidly with stand age due to spatial resource partitioning, and (4) tree death was spatially randomly distributed among larger overstory trees. The study highlights the importance of spatial heterogeneity for the long-term coexistence of shade-intolerant pioneer Douglas-fir and shade-tolerant western hemlock and western redcedar.     

灰飞虱与褐飞虱种内和种间密度效应比较

在人工气候室内比较研究了灰飞虱和褐飞虱在汕优63和秀水11上的种内和种间密度效应对其主要生物学参数的影响,结果表明两种稻飞虱均存在显著的种内竞争,主要表现为随着密度增加,若虫发育历期延长、若虫羽化率下降、雌成虫寿命缩短、每雌产卵量下降。种内密度效应与稻飞虱种类和寄主品种有显著互作关系,灰飞虱种内竞争较褐飞虱明显,二者在适宜寄主上种内竞争更为明显。同时,两种稻飞虱存在着显著的种间密度效应,主要表现为异种存在时的促进作用,即异种共存时若虫历期缩短、若虫羽化率提高、雌虫寿命延长、每雌产卵量增加。种间密度效应也与稻飞虱种类和寄主品种有显著互作关系,两种共存对灰飞虱的有利作用显著大于对褐飞虱的有利作用,在欠适宜寄主上种间互利效应更为明显。最后,本文对稻飞虱种内和种间密度效应的机制和进化意义进行了讨论。     

The arrangement of resources in patchy landscapes: effects on distribution, survival, and resource acquisition of chironomids

The spatial arrangement of resources in patchy habitats influences the distribution of individuals and their ability to acquire resources. We used Chironomus riparius, a ubiquitous aquatic insect that uses leaf particles as an important resource, to ask how the dispersion of resource patches influences the distribution and resource acquisition of mobile individuals in patchy landscapes. Two experiments were conducted in replicated laboratory landscapes (38×38 cm) created by arranging sand and leaf patches in a 5×5 grid so that the leaf patches were either aggregated or uniformly dispersed in the grid. One-day-old C. riparius larvae were introduced into the landscapes in one of three densities (low, medium, high). In experiment 1, we sampled larvae and pupae by coring each patch in each landscape 3, 6, 12, or 24 days after adding larvae. In experiment 2, emerging adults were collected daily for 42 days from each patch in each landscape. In aggregated landscapes, individuals were aggregated in one patch type or the other during a particular developmental stage, but the ”preferred” type changed depending on developmental stage and initial density. Adult emergence was lower by about 30% in all aggregated landscapes. In dispersed landscapes, individuals used both types of patch throughout their life cycles at all initial densities. Thus, patch arrangement influences the distribution of mobile individuals in landscapes, and it influences resource acquisition even when average resource abundance is identical among landscapes. Regardless of patch arrangement, high initial density caused accumulation of early instars in edge patches, 75% mortality of early instars, a 25% increase in development time, and a 60% reduction in adult emergence. Because mortality was extremely high among early-instar larvae in high-density treatments, we do not have direct evidence that the mechanism by which patch arrangement operates is density dependent. However, the results of our experiments strongly suggest that dispersion of resource patches across a landscape reduces local densities by making non-resource patches available for use, thereby reducing intraspecific competition. Received: 20 July 1999 / Accepted: 28 January 2000     

Aggregation, intraguild interactions and the coexistence of competitors on small ephemeral patches

It is well established that intraspecific aggregation has the potential to promote coexistence in communities of species competing for patchy ephemeral resources. We developed a simulation model to explore the influence of aggregation on coexistence in such communities when an important assumption of previous studies – that interspecific interactions have only negative effects on the species involved – is relaxed. The model describes a community of competing insect larvae in which an interaction that is equivalent to intraguild predation (IGP) can occur, and is unusual in that it considers species exploiting very small resource patches (carrying capacity=1). Model simulations show that, in the absence of any intraspecific aggregation, variation between species in the way that resource heterogeneity affects survival increases the likelihood of species coexistence. Simulations also show that intraspecific aggregation of the dominant competitor's eggs across resource patches can promote coexistence by reducing the importance of interspecific competition relative to that of intraspecific competition. Crucially, however, this effect is altered if one competitor indulges in IGP. In general, coexistence is only possible when the species that is capable of IGP is less effective at exploiting the shared resource than its competitor. Because it reduces the relative importance of interspecific interactions, intraspecific aggregation of the eggs of a species that is the victim of IGP actually reduces the likelihood of coexistence in parts of parameter space in which the persistence of the other species is dependent on its ability to exploit its competitor. Since resource heterogeneity, intraspecific aggregation and IGP are all common phenomena, these findings shed light on mechanisms that are likely to influence diversity in communities exploiting patchy resources.     

An evolutionary explanation of the aggregation model of species coexistence

In ecology, the 'aggregation model of coexistence' provides a powerful concept to explain the unexpectedly high species richness of insects on ephemeral resources like dung pats, fruits, etc. It suggests that females aggregate their eggs across resource patches, which leads to an increased intraspecific competition within occupied patches and a relatively large number of patches that remain unoccupied. This provides competitor-free patches for heterospecifics, facilitating species coexistence. At first glance, deliberately causing competition among the females' own offspring and leaving resources to heterospecific competitors seems altruistic and incompatible with individual fitness maximization, raising the question of how natural selection operates in favour of egg aggregation on ephemeral resource patches. Allee effects that lead to fitness maxima at intermediate egg densities have been suggested, but not yet detected. Using drosophilid flies on decaying fruits as a study system, we demonstrate a hump-shaped relationship between egg density and individual survival probability, with maximum survivorship at intermediate densities. This pattern clearly selects for egg aggregation and resolves the possible conflict between the ecological concept of species coexistence on ephemeral resources and evolutionary theory.     

Sexual Hieracium pilosella plants are better inter-specific,while apomictic plants are better intra-specific competitors

Apomixis, asexual reproduction through seeds, occurs in over 40 plant families. This widespread phenomenon can lead to the fixation of successful genotypes, resulting in a fitness advantage. On the other hand, apomicts are expected to lose their fitness advantage if the environment changes because of their limited evolutionary potential, which is due to low genetic variability and the potential accumulation of deleterious somatic mutations. Nonetheless, some apomicts have been extremely successful, for example certain apomictic accessions of Hieracium pilosella L. from New Zealand, where the plant is invasive. Here, we investigate whether the success of these apomictic accessions could be due to a fitness advantage by comparing the vegetative competitiveness of apomictic H. pilosella from New Zealand with sexual accessions of H. pilosella from Europe. Sexual and apomictic plants were grown either (A) alone (no competition), (B) in competition with the other type (intra-specific competition), (C) in competition with the grass Bromus erectus (inter-specific competition), and (D) in competition with the other type and the grass B. erectus (intra- and inter-specific competition). To distinguish effects of apomixis and the region of origin, different H. pilosella lineages were compared. Furthermore, experiments were carried out to investigate effects of the ploidy level. We show that sexual plants are better inter-specific competitors than apomicts in terms of vegetative reproduction (number of stolons) and vegetative spread (stolon length), while apomicts do better than sexuals in intra-specific competition. The magnitude of the effect was in some cases dependent on the ploidy levels of the plants. Furthermore, apomicts always produced more stolons than sexuals, suggesting potential displacement of sexuals by apomicts where they co-occur.     

Larval aggregation and competition for food in experimental populations of Chrysomya putoria (Wied.) and Cochliomyia macellaria (F.) (Dipt., Calliphoridae)

In blowflies, larval aggregation in patches of food can be both intra- and interspecific, depending upon the degree to which competitors are clumped among the patches. In the present study, the implications of spatial aggregation for larval competition was investigated in experimental populations of the introduced blowfly Chrysomya putoria and the native Cochliomyia macellaria , using data from survival to adulthood in a range of single- and double-species larval cultures. The reduction in C. macellaria survival rate in the presence of C. putoria suggests that the former species is the inferior competitor. The results on survival to adulthood for both species in single- and double-species cultures can be explained in the light of the relationship between the level of intra- and interspecific aggregation and the efficiency of the larval feeding process. The possible implications of these results for the population biology of both species in natural environments are discussed.     

Modelling Inter- and Intra-specific Competition in Loblolly Pine (Pinus taeda L.) Plantations on Cutover, Site-prepared Lands

The presence of volunteer hardwood species in loblolly pineplantations demands studies on both intra- and inter-specificcompetition in order to build growth and yield models and toguide vegetation management for these stands. This paper, basedon analyses of data collected from a thinning study, reportsan investigation of responses of loblolly pine and hardwoodspecies towards intra- and inter-specific competition. Underhigh levels of overall competition, hardwood species were morecompetitive, both intra- and inter-specifically. Intra-specificcompetition was more effective in reducing hardwood basal areagrowth than inter-specific competition under high levels ofoverall competition. However, under low levels of overall competition,intra- and inter-specific competition were quantitatively similartoward loblolly pine basal area growth. The basal area growthof hardwoods was significantly related to levels of inter-specificcompetition, but not with intra-specific effects under low levelsof overall competition. More than half of the variability inloblolly pine basal area growth under unthinned control andlight thin treatments could be accounted for by competitioneffects. However, only one third of the variability could beexplained for loblolly pine under heavy thin treatment. Forhardwood species, the percentage of growth variation accountedfor by competition was about 45%, and did not change among thethinning treatments. Different resource demands between thetwo categories of tree species and a certain amount of thinningshock were suggested by the modelling results.Copyright 1994,1999 Academic Press Neighbourhood approach, competition index, hardwood, no-interaction model, basal area, reciprocal yield law     

Genetic structure of natural Nasonia vitripennis populations: validating assumptions of sex-ratio theory

The parasitic wasp Nasonia vitripennis has been used extensively in sex allocation research. Although laboratory experiments have largely confirmed predictions of local mate competition (LMC) theory, the underlying assumptions of LMC models have hardly been explored in nature. We genotyped over 3500 individuals from two distant locations (in the Netherlands and Germany) at four polymorphic microsatellite loci to validate key assumptions of LMC theory, in terms of both the original models and more recent extensions to them. We estimated the number of females contributing eggs to patches of hosts and the clutch sizes as well as sex ratios produced by individual foundresses. In addition, we evaluated the level of inbreeding and population differentiation. Foundress numbers ranged from 1 to 7 (average 3.0 ± 0.46 SE). Foundresses were randomly distributed across the patches and across hosts within patches, with few parasitizing more than one patch. Of the hosts, 40% were parasitized by more than one foundress. Clutch sizes of individual foundresses (average 9.99 ± 0.51 SE) varied considerably between hosts. The time period during which offspring continued to emerge from a patch or host correlated strongly with foundress number, indicating that sequential rather than simultaneous parasitism is the more common. Genetic differentiation at the regional level between Germany and the Netherlands, as estimated by Slatkin's private allele method (0.11) and Hedrick's corrected G' _LT (0.23), indicates significant substructuring between regions. The level of population inbreeding for the two localities ( F _IL = 0.168) fitted the expectation based on the average foundress number per patch.     

Spatial distributions and clutch sizes of Drosophila species ovipositing on cherry fruits of different stages

In the aggregation theory, aggregation of eggs is one of important conditions for the coexistence of species. However, aggregation of eggs by clutch laying does not always promote coexistence, whereas aggregation of eggs by aggregated distributions of ovipositing females always has a significant contribution to the coexistence. In this study, spatial distributions of three Drosophila species across naturally occurring cherry fruits were studied with relation to their clutch sizes. Drosophila suzukii oviposited eggs mainly on fresh fruits on trees, and its eggs were randomly distributed across cherry fruits. The emergence data also indicated random spatial distributions of this species. Random egg distributions of this species are explained by random visits of females to fruits and the production of clutches of mostly single eggs. On the other hand, D. lutescens and D. rufa oviposited on fallen fruits, showed aggregated distributions in the emergence data, and frequently produced clutches of a few eggs. In these species, the degree of aggregation was usually significantly lower than the expectation based on random visits of females to fruits and their clutch sizes observed in the present experiments, indicating that their aggregation is unlikely to arise from aggregated distributions of ovipositing females. Thus, the spatial aggregation of these species does not necessarily lead to their coexistence.     

Aggregative Root Placement: A Feature During Interspecific Competition in Inland Sand-Dune Habitats

Segregation of roots is frequently observed in competing root systems. However, recently, intensified root growth in response to a neighbouring plant has been described in pot experiments [Gersani M, Brown J S, O'Brien E E, Maina G M and Abramsky Z 2001. J. Ecol. 89, 660–669]. This paper examines whether intense root growth towards a neighbour (aggregation) plays a role in competitive interactions between plant species from open nutrient-poor mid-European sand ecosystems. In a controlled field-competition experiment, root distribution patterns of intra- and interspecific pairs as well as single control plants of Corynephorus canescens, Festuca psammophila, Hieracium pilosella, Hypochoeris radicata and Conyza canadensis were investigated after one growing season. Under intraspecific competition plants tended to segregate their root systems, while under interspecific competition most species tended to aggregate roots towards their neighbours even at the expense of root development at the opposite competition-free side of the target. Preference of a root aggregation strategy over the occupation of competition-free soil in interspecific competition emphasizes the importance of contesting between individuals in relation to mere resource acquisition. It is suggested that in the presence of a competitor the plants might use root aggregation as a defensive reaction to maintain a strong competitive response and exclusive access to the resources of already occupied soil volumes.     

苔藓植物相互作用的研究进展

综述了苔藓植物种内和种间正相互作用、种内和种间竞争、生态位与共存等的研究现状及其进展.苔藓植物常为严酷环境中的优势植物,并在保持水分和获取光及养分资源两方面存在权衡.因严酷环境中竞争的重要性相对较低,苔藓植物的正相互作用较为多见.然而,在苔藓植物种内、种间及其与维管植物之间,均不乏竞争的实例.在严酷性较低的环境如过湿的泥炭地中,竞争的重要性增加.多种苔藓植物间可存在竞争等级,该等级往往随环境的变化而变化.随机过程、物种的殖民本质以及更新策略的差异可导致苔藓植物生态位的重叠与共存,但竞争导致生态位分化也是苔藓植物共存的机理之一.苔藓植物不应简单地归类于耐压-杂草型生活史策略者,竞争仍是构建某些苔藓植物群落及其与维管植物共存群落的重要因子之一.     

Habitat-specific estimates of competition in stream salmonids: A field test of the isodar model of habitat selection

Summary The population densities of sympatric Atlantic salmon,Salmo salar and brook charr,Salvelinus fontinalis, were measured in riffle and pool stream habitats to test whether non-linear isodars, a multispecific model of habitat selection based on ideal distribution assumptions, could (1) predict the distribution of densities between habitats and (2) reproduce the processes postulated to underlie spatial segregation and species interactions in previous laboratory and field studies. The model provided a good fit to observed density patterns and indicated that habitat suitability declined non-linearly with increased heterospecific competitor densities. Competitive effects in riffles appeared to be due to exploitative resource use, with salmon always emerging as the superior competitor. No evidence was found for interference competition in riffles. In contrast, interspecific competition in pools seemed to occur through exploitation and interference. The specific identity of the superior competitor in pools depended on the density of both species; pools provided the charr with refuge from competition with the salmon, presumably through the adoption by the charr of density-dependent behaviours, such as schooling and group foraging, that mitigated the negative impact of the salmon. Charr were displaced from the riffles toward the pools as the total salmon density increased. The isodar analysis, based on limited density data, successfully reproduced the processes suggested to underlie spatial segregation in previous field and laboratory studies and provided new insights into how changes in competitor densities modify habitat suitability in this system.     

闽东北海域假长缝拟对虾种群分布格局强度

以负二项参数、Mrisita指数、平均拥挤度、聚块指数和扩散系数为种群分布格局强度指标,分析了闽东北海域假长缝拟对虾(Parapenaeus fissuroides Crosnier)种群分布格局强度,探讨了种群分布格局强度生态学意义及其影响因子。结果显示:假长缝拟对虾种群聚集强度和聚块性夏季最强,其余依次为秋季、春季和冬季,平均拥挤度夏季最大,其次是秋季、冬季,春季最小,扩散程度春季最大,其次是冬季、秋季,夏季最小。负二项参数和Mrisita指数表征种群聚集强度,聚块指数体现种群斑块的组成形式,个体平均拥挤度指数反映平均个体实际的拥挤程度,扩散系数反映种群扩散程度。饵料浮游动物是影响假长缝拟对虾种群聚集强度的主要因子,种内竞争是影响假长缝拟对虾种群扩散程度的主要因子。     

Effects of spatial aggregation on competition,complementarity and resource use

Abstract The spatial distributions of most species are aggregated to varying degrees. A limited number of studies have examined the effects of spatial aggregation on interspecific and intraspecific interactions, generally finding that spatial aggregation can enhance coexistence between species by reducing the capacity for interspecific competition. Less well studied are the effects of spatial aggregation on complementarity (i.e. differences in resource use strategies) and resource use. Our primary hypothesis was that spatial aggregation reduces the complementarity between species owing to: (i) less interspecific interactions as a result of spatial separation; and (ii) less differences between species as a result of phenotypic plasticity. We further postulate that these negative effects of spatial aggregation on complementarity will reduce resource use by the community. Here we test these hypotheses in a pot experiment in which we applied three levels of spatial aggregation to three sets of two‐species mixtures of herbaceous perennial plant species from native grasslands of south‐eastern Australia. Both root and shoot biomass were significantly affected by spatial aggregation, although the nature of these affects depended upon the species involved, and the relative strengths of interspecific versus intraspecific competition. Complementarity between species in the distribution of their green leaves decreased significantly as spatial aggregation increased for one of the species mixtures, providing some evidence in support of our hypothesis that aggregation reduces complementarity through phenotypic plasticity. Spatial aggregation also altered light interception and use of soil moisture resources, although these effects were dependent on the species involved. We suggest that clear effects of spatial aggregation on complementarity and resource use may be obscured by the idiosyncratic way in which neighbour identity influences plant growth and hence plant size, limiting the ability to generalize, at the community level, any underlying effects of spatial pattern on ecological process.     

Clutch-size behavior and coexistence in ephemeral-patch competition models

Systems of patchy, ephemeral resources often support surprisingly diverse assemblages of consumer insects. Aggregation of consumer individuals over the landscape of patches has been suggested as one mechanism that can stabilize competition among consumer species. One mechanism for larval aggregation is the laying of eggs in clutches by females traveling among patches to distribute their total fecundity. We use simulation models to explore the consequences, for coexistence of competitors, of larval aggregation that arises from clutch laying. Contrary to some previous treatments, we find that clutch laying can be strongly stabilizing and under certain conditions can be sufficient to allow competitors to coexist stably. We extend these models by considering clutch size as a variable that responds to the abundance of resource patches. Such a relationship might be expected because females should lay their eggs in fewer but larger clutches when the cost of travel among patches is high (because patches are rare). When females adjust clutch size in response to resource abundance, coexistence can be easiest when resource patches are scarce and most difficult when resources are abundant.     

Remating and sperm displacement in a natural population of Drosophila buzzatii inferred from mother-offspring analysis of microsatellite loci

Prospects for estimation of parameters of models of sperm competition from field data have improved recently with the development of methods that employ multilocus genotype data from brood-structured samples. Sperm competition in Drosophila buzzatii is of special interest because it is possible to directly observe the breeding behaviour of this species in its natural habitat of rotting cactus. Previous laboratory experiments showed that this species exhibits an unusual pattern of frequent remating and sperm partitioning. This paper reports the first attempt to estimate the frequency of female remating and sperm competition in natural populations of D. buzzatii. For the Australian population studied, the mean remating frequency was lower (alpha = 2.12-2.20) than previously estimated in laboratory experiments with the same population, whereas mean sperm displacement (beta = 0.69-0.71) fell within the limits of previous laboratory results. The evolution of the D. buzzatii mating system is discussed.     

The genetic effects of competition in seaweed flies

In spite of abundant evidence that intra- and inter-specific competition occurs in natural communities, there is surprisingly little to suggest it is a major force promoting genetic change. This report assesses the genetic effects of competition in two species of seaweed fly, Coelopa frigida and C. pilipes. In laboratory cultures of C. frigida the relative survival of heterozygotes at the Adh locus, which was being used as a marker for the large αβ chromosomal inversion, was greater than that of homozygotes. In monocultures of C. frigida this competitive superiority was dependent on larval density. At low densities facilitation was seen, whereas at high larval densities there was competition. In mixed cultures of the two species, interspecific competition contributed to the differential mortality of C. frigida , and observations of natural populations suggested that competition may have similar effects to those described in laboratory culture. A possible mechanism involving the supply of nutritive microorganisms is proposed to underly both intra- and inter-specific competition. In seaweed flies, competition and the consequent differential mortality appear to be forces maintaining rather than reducing genetic variation.     

Body size mediated coexistence of consumers competing for resources in space

Body size is a major phenotypic trait of individuals that commonly differentiates co-occurring species. We analyzed inter-specific competitive interactions between a large consumer and smaller competitors, whose energetics, selection and giving-up behaviour on identical resource patches scaled with individual body size. The aim was to investigate whether pure metabolic constraints on patch behaviour of vagile species can determine coexistence conditions consistent with existing theoretical and experimental evidence. We used an individual-based spatially explicit simulation model at a spatial scale defined by the home range of the large consumer, which was assumed to be parthenogenic and semelparous. Under exploitative conditions, competitive coexistence occurred in a range of body size ratios between 2 and 10. Asymmetrical competition and the mechanism underlying asymmetry, determined by the scaling of energetics and patch behaviour with consumer body size, were the proximate determinant of inter-specific coexistence. The small consumer exploited patches more efficiently, but searched for profitable patches less effectively than the larger competitor. Therefore, body-size related constraints induced niche partitioning, allowing competitive coexistence within a set of conditions where the large consumer maintained control over the small consumer and resource dynamics. The model summarises and extends the existing evidence of species coexistence on a limiting resource, and provides a mechanistic explanation for decoding the size-abundance distribution patterns commonly observed at guild and community levels.