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.     

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.     

Effects of nematode parasitism on mycophagous drosophilid species in northern Japan

Mycophagous drosophilids and their nematode parasites were studied in an intensive census conducted throughout the active season over four years (2000–2003) in Hokkaido, northern Japan. Using census data, I assessed the prevalence of nematode parasitism through seasons, determined its effects on drosophilid female fertility and tested the “disproportionate parasitism hypothesis” to evaluate its effects on drosophilid community structure. Over the four‐year census, a total of 43 fungal species were found on the census route and 18 177 adult drosophilid flies were collected. The yearly parasitism rate on the total drosophilid community varied from 6.2% in 2000 to 5.1% in 2001 and 3.0% in 2002 and 2003. There was strong seasonal variation in the prevalence of parasitism. In addition, parasitism rates varied among drosophilid species. All parasitized females carried only small numbers of eggs. Thus, nematode parasitism had a strong deleterious effect on the fitness of infected flies. The relative abundance of drosophilid species and parasitism rate were not significantly correlated. Thus, the disproportionate parasitism hypothesis likely does not apply to the mycophagous drosophilid community in Japan.     

Effects of parasitoids on a mycophagous drosophilid community in northern Japan and an evaluation of the disproportionate parasitism hypothesis

In a host–parasitoid system comprising mycophagous drosophilids and their parasitoids, the drosophilid and parasitoid species assemblages, host use, and the prevalence of parasitism were assessed, and the “disproportionate parasitism hypothesis” was examined with consideration given to yearly variations. The mycophagous drosophilids, their fungal food resources and parasitoids were studied by carrying out an intensive census throughout the activity seasons of 4 years (2000–2003) in Hokkaido, northern Japan. Five hymenopterous parasitoid species, four braconids and one eucoilid, were found. Parasitoids of mycophagous drosophilids are reported for the first time from Asia. Most parasitism (99.2%) was by braconids, in contrast to the dominance of eucoilids in Europe. Parasitism was restricted to the summer, and the rate was high from early July to early August every year. There was considerable yearly variation in the composition of abundant fungus, drosophilid and parasitoid species, especially between 2000 and 2001. The alternation of dominant host species was coupled with the alternation of dominant parasitoid species that differed in host use. Despite the yearly variation in the system, the most dominant host species suffered disproportionately heavy parasitism by the correspondingly dominant parasitoid species every year. The parasitism rate was positively correlated with the relative host abundance. This thus indicates that the disproportionate parasitism mechanism may operate, via which species coexistence is promoted by a higher rate of parasitism of the dominant species.     

Taxonomic scale-dependence of habitat niche partitioning and biotic neighbourhood on survival of tropical tree seedlings

In order to differentiate between mechanisms of species coexistence, we examined the relative importance of local biotic neighbourhood, abiotic habitat factors and species differences as factors influencing the survival of 2330 spatially mapped tropical tree seedlings of 15 species of Myristicaceae in two separate analyses in which individuals were identified first to species and then to genus. Using likelihood methods, we selected the most parsimonious candidate models as predictors of 3 year seedling survival in both sets of analyses. We found evidence for differential effects of abiotic niche and neighbourhood processes on individual survival between analyses at the genus and species levels. Niche partitioning (defined as an interaction of taxonomic identity and abiotic neighbourhood) was significant in analyses at the genus level, but did not differentiate among species in models of individual seedling survival. By contrast, conspecific and congeneric seedling and adult density were retained in the minimum adequate models of seedling survival at species and genus levels, respectively. We conclude that abiotic niche effects express differences in seedling survival among genera but not among species, and that, within genera, community and/or local variation in adult and seedling abundance drives variation in seedling survival. These data suggest that different mechanisms of coexistence among tropical tree taxa may function at different taxonomic or phylogenetic scales. This perspective helps to reconcile perceived differences of importance in the various non-mutually exclusive mechanisms of species coexistence in hyper-diverse tropical forests.     

Higher species diversity explained by stronger spatial aggregation across six neotropical Drosophila communities

Spatial aggregation of competitors over resource patches is generally accepted as an important mechanism maintaining coexistence of species in insect communities exploiting fragmented resources. However, its quantitative effects on local diversity, i.e. the relationship between the degree of aggregation in a community and community diversity, remain unexplored. In this paper, we tested whether stronger spatial aggregation does lead to the predicted higher local diversity. We compared six species-rich Drosophila communities exploiting decaying fruits in central Panama, monitored over one full year (> 25 generations). We found a clear positive relationship between the overall degree of aggregation and community diversity. In addition, aggregation over fruit trees was found to contribute greatly to the overall degree of aggregation and was largely responsible for the observed relationship between aggregation and diversity across communities. In addition, both diversity and aggregation strength were lower in communities in disturbed habitats, which was explained by altered spatial distribution of fruiting trees. This study shows that the aggregation model cannot only explain coexistence, but also differences in local diversity.     

Strong plant-soil associations in a heterogeneous subtropical broad-leaved forest

The relative importance of deterministic and neutral processes on community assembly is currently a topic of much debate among ecologists. Analyzing species-environment associations is an effective way to assess the importance of deterministic process such as niche differentiation, but both habitat association and dispersal limitation can produce similar patterns of spatial aggregation in species. Therefore, it is crucial to control for the impact of dispersal limitation on species distributions when analyzing species-environment associations. We sampled soil with high resolutions in a 24 ha stem-mapped subtropical forest and tested plant-soil associations. We controlled for the influence of dispersal limitation by employing the homogeneous Thomas process to simulate the effect of dispersal limitation on the aggregation of tree species. After controlling for the effect of dispersal limitation, we found that the spatial heterogeneity of soil properties was associated with distributions of 88.2% (90 of 102 species) of tree species in this subtropical forest. Furthermore, not only did soil properties influence the distribution of tree species, but also tree species tended to affect properties of the soil around them. The soil factors most strongly influencing species distributions were TC, TN, TP, K, Mg, Si, soil moisture, and bulk density. We found the spatial heterogeneity of soil properties to be strongly associated with tree species distributions. Niche partitioning of soil gradients contributed substantially to species coexistence in this subtropical forest.     

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.     

Coexistence via resource partitioning fails to generate an increase in community function

Classic ecological theory suggests that resource partitioning facilitates the coexistence of species by reducing inter-specific competition. A byproduct of this process is an increase in overall community function, because a greater spectrum of resources can be used. In contrast, coexistence facilitated by neutral mechanisms is not expected to increase function. We studied coexistence in laboratory microcosms of the bactivorous ciliates Paramecium aurelia and Colpidium striatum to understand the relationship between function and coexistence mechanism. We quantified population and community-level function (biomass and oxygen consumption), competitive interactions, and resource partitioning. The two ciliates partitioned their bacterial resource along a size axis, with the larger ciliate consuming larger bacteria than the smaller ciliate. Despite this, there was no gain in function at the community level for either biomass or oxygen consumption, and competitive effects were symmetrical within and between species. Because other potential coexistence mechanisms can be ruled out, it is likely that inter-specific interference competition diminished the expected gain in function generated by resource partitioning, leading to a system that appeared competitively neutral even when structured by niche partitioning. We also analyzed several previous studies where two species of protists coexisted and found that the two-species communities showed a broad range of biomass levels relative to the single-species states.     

Habitat filtering drives the local distribution of congeneric species in a Brazilian white‐sand flooded tropical forest

The investigation of ecological processes that maintain species coexistence is revealing in naturally disturbed environments such as the white‐sand tropical forest, which is subject to periodic flooding that might pose strong habitat filtering to tree species. Congeneric species are a good model to investigate the relative importance of ecological processes that maintain high species diversity because they tend to exploit the same limiting resources and/or have similar tolerance limits to the same environmental conditions due to their close phylogenetic relationship. We aim to find evidence for the action and relative importance of different processes hypothesized to maintain species coexistence in a white‐sand flooded forest in Brazil, taking advantage of data on the detailed spatial structure of populations of congeneric species. Individuals of three Myrcia species were tagged, mapped, and measured for diameter at soil height in a 1‐ha plot. We also sampled seven environmental variables in the plot. We employed several spatial point process models to investigate the possible action of habitat filtering, interspecific competition, and dispersal limitation. Habitat filtering was the most important process driving the local distribution of the three Myrcia species, as they showed associations, albeit of different strength, to environmental variables related to flooding. We did not detect spatial patterns, such as spatial segregation and smaller size of nearby neighbors, that would be consistent with interspecific competition among the three congeneric species and other co‐occurring species. Even though congeners were spatially independent, they responded to differences in the environment. Last, dispersal limitation only led to spatial associations of different size classes for one of the species. Given that white‐sand flooded forests are highly threatened in Brazil, the preservation of their different habitats is of utmost importance to the maintenance of high species richness, as flooding drives the distribution of species in the community.     

Do ectotherms partition thermal resources? We still do not know

Partitioning of the niche space is a mechanism used to explain the coexistence of similar species. Ectotherms have variable body temperatures and their body temperatures influence performance and, ultimately, fitness. Therefore, many ectotherms use behavioral thermoregulation to avoid reduced capacities associated with body temperatures far from the optimal temperature for performance. Several authors have proposed that thermal niche partitioning in response to interspecific competition is a mechanism that allows the coexistence of similar species of ectotherms. We reviewed studies on thermal resource partitioning to evaluate the evidence for this hypothesis. In almost all studies, there was insufficient evidence to conclude unequivocally that thermal resource partitioning allowed species coexistence. Future studies should include sites where species are sympatric and sites where they are allopatric to rule out alternative mechanisms that cause differences in thermal traits between coexisting species. There is evidence of conservatism in the evolution of most thermal traits across a wide range of taxa, but thermal performance curves and preferred temperatures do respond to strong selection under laboratory conditions. Thus, there is potential for selection to act on thermal traits in response to interspecific competition. Nevertheless, more stringent tests of the thermal resource partitioning hypothesis are required before we can assess whether it is widespread in communities of ectotherms in nature.     

Predictability, toxicity, and trophic niche breadth in fungus-feeding Drosophilidae (Diptera)

Abstract. 1. The mycophagous Drosophilidae flies in eastern North America display a range of ecological responses to their fungal trophic resources. Some ( Drosophilu duncani Sturtevant, and two species labelled as Mycodrosophila cluytonae Wheeler and Takada) specialize on the bracket fungi of the family Polyporaceae. Other species ( Drosophila falleni Wheeler, D.recens Wheeler, D.putridu Sturtevant and D.tripunctuta Loew) are broad generalists, non-selectively consuming a diverse array of Basidioniycete mushrooms. D.restaceu von Roser and D.ordinuria Coquillett utilize a broad subset of the Basidio-mycetes, while M.dimidiata hew feeds on some Basidioniycetes, including a variety of Clavariaceae (coral fungi) and Heterobasidiomycetes (jelly fungi) not commonly used by other flies, and Ascomycetes (cup fungi).
2. Unlike some phytophagous insects, host chemistry seems to have little effect on host preferences. The mycophagous drosophilids do not avoid fungi thought to be toxic or undigestible to other Diptera.
3. Resource predictability appears to delimit trophic niches in the mycophagous guild. The average duration of individual host mushroom caps is negatively correlated with three measures of trophic niche breadth. Specialist fly species utilize long-lasting resources, while more generalized mycophagous species include progressively ephemeral fungi in their diets.     

Scaling up from local competition to regional coexistence across two scales of spatial heterogeneity: insect larvae in the fruits of Apeiba membranacea

Species that live in patchy and ephemeral habitats can compete strongly for resources within patches at a small scale. The ramifications of these interactions for population dynamics and coexistence at regional scales will depend on the intraspecific and interspecific distributions of individuals among patches. Spatial heterogeneity due to independent aggregation of competitors among patchy habitats is an important mechanism maintaining species diversity. I describe regional patterns of aggregation for four species of insect larvae in the fruits of Apeiba membranacea, a Neotropical rainforest tree. This aggregation results from variation in densities at a small scale (among the fruits under a single tree), compounded by significant variation among trees in both mean densities and degrees of aggregation. Both the degrees of aggregation and mean densities are statistically independent within and across species at both spatial scales. I evaluate the regional consequences of these spatial patterns by using maximum likelihood methods to parameterize a model that includes both explicit measures of the strength of competition and spatial variation at both within- and among-tree spatial scales. Despite strong competitive interactions among these species, during 2 years the observed spatial variation at both scales combined was sufficient to explain the coexistence of these species, although other coexistence mechanisms may also operate simultaneously. The observed spatial variation at small spatial scales may not be sufficient for coexistence, indicating the importance of considering multiple sources of spatial heterogeneity when scaling up from experiments that investigate local interactions to regional patterns of coexistence.     

Coexistence of drosophilid flies: Aggregation, patch size diversity and parasitism

We carried out field experiments to investigate the coexistence of Drosophila species in domestic and forest areas on the basis of the aggregation model. Three cosmopolitan species Drosophila simulans Sturtevant, Drosophila melanogaster Meigen and Drosophila immigrans Sturtevant, and a native species, Drosophila auraria Peng, emerged abundantly from banana placed at the domestic station, while Drosophila immigrans and five native species, Drosophila lutescens Okada, Drosophila rufa Kikkawa and Peng, Drosophila bizonata Kikkawa and Peng, Drosophila sternopleuralis Okada and Kurokawa and Scaptodrosophila coracina (Kikkawa and Peng), were abundant at the forest station. The present analysis suggests that their coexistence was facilitated by the aggregation mechanism. In the cosmopolitan species, the density of individuals that emerged from patches increased with the increase of patch size, but the relationship between fly density and patch size was not clear in the native species. This difference in distribution patterns between the cosmopolitan and native species is likely to be due to the difference in the female visiting behavior. In the present analysis, however, it was not clear whether patch size diversity facilitated their coexistence or not. The effect of patch size diversity may have been masked, because the effect of aggregation was more prominent. The rate of parasitism by wasps was high in October at the domestic station, and in May and June at the forest station. The present result suggests that the rate of parasitism was density-dependent, at least at the domestic station, and therefore parasitism facilitates the coexistence of drosophilid species in domestic areas.     

Scaling up from local competition to regional coexistence across two scales of spatial heterogeneity: insect larvae in the fruits of <Emphasis Type="Italic">Apeiba membranacea</Emphasis>

Species that live in patchy and ephemeral habitats can compete strongly for resources within patches at a small scale. The ramifications of these interactions for population dynamics and coexistence at regional scales will depend on the intraspecific and interspecific distributions of individuals among patches. Spatial heterogeneity due to independent aggregation of competitors among patchy habitats is an important mechanism maintaining species diversity. I describe regional patterns of aggregation for four species of insect larvae in the fruits of Apeiba membranacea, a Neotropical rainforest tree. This aggregation results from variation in densities at a small scale (among the fruits under a single tree), compounded by significant variation among trees in both mean densities and degrees of aggregation. Both the degrees of aggregation and mean densities are statistically independent within and across species at both spatial scales. I evaluate the regional consequences of these spatial patterns by using maximum likelihood methods to parameterize a model that includes both explicit measures of the strength of competition and spatial variation at both within- and among-tree spatial scales. Despite strong competitive interactions among these species, during 2 years the observed spatial variation at both scales combined was sufficient to explain the coexistence of these species, although other coexistence mechanisms may also operate simultaneously. The observed spatial variation at small spatial scales may not be sufficient for coexistence, indicating the importance of considering multiple sources of spatial heterogeneity when scaling up from experiments that investigate local interactions to regional patterns of coexistence.     

群落中克隆植物的重要性

综述了群落水平上克隆植物的重要性,克隆植物的生态习性,克隆植物的竞争关系等方面的研究进展,并试图在克隆植物竞争关系的背景下,结合其生境状况,探讨植物克隆性与植物物种多样性的关系。克隆植物的等级系统（基株—分株系统—分株）使其具有克隆性所赋予的多样的生活史、资源利用以及空间占据方式。列举了克隆植物的特性及其在群落中的作用。这些例子表明,克隆性强烈地影响和制约着植物群落的空间格局与竞争关系。然而克隆性也能通过权衡活动性与局部持久性来缓解对物种共存的抑制。克隆植物具有在时空尺度上分株间相互调节的机制,直接体现在群落中小尺度（个体与个体间）与大尺度（种群与种群间）间的相互作用上。丰富了传统的竞争和生态位划分理论,为群落中物种共存提供了合理解释。因此克隆植物在群落中的出现拓宽了群落系统潜在机制的范围。     

Drivers of symbiont diversity in freshwater snails: a comparative analysis of resource availability,community heterogeneity,and colonization opportunities

Decades of community ecology research have highlighted the importance of resource availability, habitat heterogeneity, and colonization opportunities in driving biodiversity. Less clear, however, is whether a similar suite of factors explains the diversity of symbionts. Here, we used a hierarchical dataset involving 12,712 freshwater snail hosts representing five species to test the relative importance of potential factors in driving symbiont richness. Specifically, we used model selection to assess the explanatory power of variables related to host species identity, resource availability (average body size, host density), ecological heterogeneity (richness of hosts and other taxa), and colonization opportunities (wetland size and amount of neighboring wetland area) on symbiont richness in 146 snail host populations in California, USA. We encountered a total of 23 taxa of symbionts, including both obligatory parasites such as digenetic trematodes as well as more commensal, mutualistic, or opportunistic groups such as aquatic insect larvae, annelids, and leeches. After validating richness estimates per host population using species accumulative curves, we detected positive effects on symbiont richness from host body size, total richness of the aquatic community, and colonization opportunities. Neither snail density nor the richness of snail species accounted for significant variation in symbiont diversity. Host species identity also affected symbiont richness, with higher gamma and average alpha diversity among more common host species with higher local abundances. These findings highlight the importance of multiple, concurrent factors in driving symbiont richness that extend beyond epidemiological measures of host abundance or host diversity alone.     

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.     

Temporal resource partitioning mitigates interspecific competition and promotes coexistence among insect parasites

A key to understanding life's great diversity is discerning how competing organisms divide limiting resources to coexist in diverse communities. While temporal resource partitioning has long been hypothesized to reduce the negative effects of interspecific competition, empirical evidence suggests that time may not often be an axis along which animal species routinely subdivide resources. Here, we present evidence to the contrary in the world's most biodiverse group of animals: insect parasites (parasitoids). Specifically, we conducted a meta-analysis of 64 studies from 41 publications to determine if temporal resource partitioning via variation in the timing of a key life-history trait, egg deposition (oviposition), mitigates interspecific competition between species pairs sharing the same insect host. When competing species were manipulated to oviposit at (or near) the same time in or on a single host in the laboratory, competition was common, and one species was typically inherently superior (i.e. survived to adulthood a greater proportion of the time). In most cases, however, the inferior competitor could gain a survivorship advantage by ovipositing earlier (or in a smaller number of cases later) into shared hosts. Moreover, this positive (or in a few cases negative) priority advantage gained by the inferior competitor increased as the interval between oviposition times became greater. The results from manipulative experiments were also correlated with patterns of life-history timing and demography in nature: the more inherently competitively inferior a species was in the laboratory, the greater the interval between oviposition times of taxa in co-occurring populations. Additionally, the larger the interval between oviposition times of competing taxa, the more abundant the inferior species was in populations where competitors were known to coexist. Overall, our findings suggest that temporal resource partitioning via variation in oviposition timing may help to facilitate species coexistence and structures diverse insect communities by altering demographic measures of species success. We argue that the lack of evidence for a more prominent role of temporal resource partitioning in promoting species coexistence may reflect taxonomic differences, with a bias towards larger-sized animals. For smaller species like parasitic insects that are specialized to attack one or a group of closely related hosts, have short adult lifespans and discrete generation times, compete directly for limited resources in small, closed arenas and have life histories constrained by host phenology, temporal resource subdivision via variation in life history may play a critical role in allowing species to coexist by alleviating the negative effects of interspecific competition.