Predator-mediated coexistence in laboratory communities of mycophagous Drosophila (Diptera: Drosophilidae)

Abstract. 1. This laboratory experiment examined the effects of interspecific competition and predation by Ontholestes cingulatus Gravenhorst (Coleoptera: Staphylinidae) on three species of mycophagous Drosophila (Diptera: Drosophilidae): D.tripunctata Loew, D.falleni Wheeler and D.putrida Sturtevant.
2. Single-species and three-species assemblages were exposed to single commercial mushrooms on wet pine shavings in 200 ml culture bottles. A predacious rove beetle (Ontholestes cingulatus) was present in half of the three-species replicates. The stocked adult flies and beetles were removed after 4 days, and the number, biomass and mean mass of emerging progeny was recorded.
3. For all three species the abundance and biomass of the progeny emerging in the 'no predator' communities' was significantly less than for the progeny emerging in single-species replicates, suggesting an interspecific competitive effect. D.tripunctata was the competitive dominant; it emerged in abundance from all seven three-species 'no predator' communities while D.putrida and D.falleni were often excluded.
4. The decrease in production was attributed to strong interspecific competition among larva and not interference among ovipositing adults.
5. Predation on ovipositing adults significantly reduced the number and biomass of D.tripunctata progeny emerging, and indirectly facilitated the number and biomass of emerging D.falleni and D.putrida.Predation on adults reduced larval recruitment, relaxed larval competition, and released the inferior competitors.     

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.     

Stochastic models for mainland-island metapopulations in static and dynamic landscapes

This paper has three primary aims: to establish an effective means for modelling mainland-island metapopulations inhabiting a dynamic landscape; to investigate the effect of immigration and dynamic changes in habitat on metapopulation patch occupancy dynamics; and to illustrate the implications of our results for decision-making and population management. We first extend the mainland-island metapopulation model of Alonso and McKane [Bull. Math. Biol. 64:913–958, 2002] to incorporate a dynamic landscape. It is shown, for both the static and the dynamic landscape models, that a suitably scaled version of the process converges to a unique deterministic model as the size of the system becomes large. We also establish that, under quite general conditions, the density of occupied patches, and the densities of suitable and occupied patches, for the respective models, have approximate normal distributions. Our results not only provide us with estimates for the means and variances that are valid at all stages in the evolution of the population, but also provide a tool for fitting the models to real metapopulations. We discuss the effect of immigration and habitat dynamics on metapopulations, showing thatmainland-like patches heavily influence metapopulation persistence, and we argue for adopting measures to increase connectivity between this large patch and the other island-like patches. We illustrate our results with specific reference to examples of populations of butterfly and the grasshopperBryodema tuberculata.     

Stochastic niche structure and diversity maintenance in the T cell repertoire

The reliability of the immune response to pathogenic challenge depends critically on the size and diversity of the T cell repertoire. We study naïve T cell repertoire diversity maintenance by a stochastic model that incorporates the concept of competition between T cells for survival stimuli emanating from self-antigen presenting cells (APCs). In the mean field approximation we show that clonotype extinction is certain and compute mean extinction times. We introduce the concept of mean niche overlap and show that clones with a mean niche overlap greater than one have a short repertoire lifespan. This selection differential induces minimal recognition commonality between T cell receptors (TCRs) resulting in a diverse T cell repertoire.     

Interacting influence of mycorrhizal symbiosis and competition on plant diversity in tallgrass prairie

In tallgrass prairie, plant species interactions regulated by their associated mycorrhizal fungi may be important forces that influence species coexistence and community structure; however, the mechanisms and magnitude of these interactions remain unknown. The objective of this study was to determine how interspecific competition, mycorrhizal symbiosis, and their interactions influence plant community structure. We conducted a factorial experiment, which incorporated manipulations of abundance of dominant competitors, Andropogon gerardii and Sorghastrum nutans, and suppression of mycorrhizal symbiosis using the fungicide benomyl under two fire regimes (annual and 4-year burn intervals). Removal of the two dominant C₄ grass species altered the community structure, increased plant species richness, diversity, and evenness, and increased abundance of subdominant graminoid and forb species. Suppression of mycorrhizal fungi resulted in smaller shifts in community structure, although plant species richness and diversity increased. Responses of individual plant species were associated with their degree of mycorrhizal responsiveness: highly mycorrhizal responsive species decreased in abundance and less mycorrhizal responsive species increased in abundance. The combination of dominant-grass removal and mycorrhizal suppression treatments interacted to increase synergistically the abundance of several species, indicating that both processes influence species interactions and community organization in tallgrass prairie. These results provide evidence that mycorrhizal fungi affect plant communities indirectly by influencing the pattern and strength of plant competitive interactions. Burning strongly influenced the outcome of these interactions, which suggests that plant species diversity in tallgrass prairie is influenced by a complex array of interacting processes, including both competition and mycorrhizal symbiosis. Received: 7 April 1999 / Accepted: 30 July 1999     

大型食草动物的聚群规则:印度西部热带干旱森林中有蹄类动物局域和区域尺度的多样性格局

竞争能够塑造自然群落并由其给出可用备择模型检验的一般性聚集规则。关于多种动物集合的竞争结构证据至今还非常少见,我们给出了一个备择模型分析用于检验印度西部热带干旱森林中当地5种大型食草类动物(花鹿Axisaxis,印度大蓝羚Boselaphustragocamelus,水鹿Cervusunicolor,印度瞪羚Gazellabennetti,野猪Susscrofa)是否符合一般性集合规则。使用蒙特卡罗模拟分析了动物的生态位重叠和身体大小比率的类似性,结果表明当地这些种的集合不是竞争结构决定的,其内部阶元结构处于亚最适种丰富度。另外2种有蹄类(印度黑羚Antilopecervicapra,四角羚羊Tetracerusquadricornis)属于地域性种库的组成部分,但缺失于这种集合;身体大小组合在地域性物种集合中不是随机的,然而在当地集合中能够通过随机性机会获得。对于观察到的类型,我们认为由于灭绝留下的统计学空缺似乎最能解释现存集合。由于天敌(大型食肉类)能够减少猎物物种的种间竞争,使得这种无规则生态位配置具有成立的可能性。由于对四角羚羊(T.quadricornis)的生物学和保护现状知之甚少,未将该种包括在该聚集中。我认为对这类较少被人类了解的物种其保护问题被忽视了,今后获得这些稀少的地方特有物种的生态学资料是一项紧急课题.     

Minireviews: Neighborhood Effects, Disturbance Severity, and Community Stability in Forests

A theoretical framework and conceptual model for temporal stability of forest tree-species composition was developed based on a synthesis of existing studies. The model pertains primarily to time periods of several tree lifetimes (several hundred to a few thousand years) at the neighborhood and stand spatial scales (0.01–10 ha), although a few extensions to the landscape scale are also made. The cusp catastrophe was chosen to illustrate compositional dynamics at the stand level for jack pine, northern hardwood, and white pine forests in the Great Lakes Region of the United States and for tropical rainforests in the northern Amazon basin. The models feature a response surface (degree of dominance by late-successional species) that depends on two variables: type of neighborhood effects of the dominant tree species and severity of disturbances. Neighborhood effects are processes that affect the chance of a species replacing itself at the time of disturbance (they can be positive, neutral, or negative) and are of two types: overstory–undestory effects, such as the presence of advanced reproduction; and disturbance-activated effects, such as serotinous seed rain. Disturbance severity is the proportion of trees killed during a disturbance. Interactions between neighborhood effects and disturbance severity can lead to either punctuated stability (dramatic but infrequent change in composition, in those forests dominated by species with positive neighborhood effects) or succession (continuous change, in those forests dominated by species with neutral-negative neighborhood effects). We propose that neighborhood effects are a major organizing factor in forest dynamics that provide a link across spatial scales between individual trees and disturbance/patch dynamics at the stand and landscape scales. Received 23 June 1998; accepted 16 December 1998.     

Disentangling the relative importance of climate,size and competition on tree growth in Iberian forests: implications for forest management under global change

Most large‐scale multispecies studies of tree growth have been conducted in tropical and cool temperate forests, whereas Mediterranean water‐limited ecosystems have received much less attention. This limits our understanding of how growth of coexisting tree species varies along environmental gradients in these forests, and the implications for species interactions and community assembly under current and future climatic conditions. Here, we quantify the absolute effect and relative importance of climate, tree size and competition as determinants of tree growth patterns in Iberian forests, and explore interspecific differences in the two components of competitive ability (competitive response and effect) along climatic and size gradients. Spatially explicit neighborhood models were developed to predict tree growth for the 15 most abundant Iberian tree species using permanent‐plot data from the Spanish Second and Third National Forest Inventory (IFN). Our neighborhood analyses showed a climatic and size effect on tree growth, but also revealed that competition from neighbors has a comparatively much larger impact on growth in Iberian forests. Moreover, the sensitivity to competition (i.e. competitive response) of target trees varied markedly along climatic gradients causing significant rank reversals in species performance, particularly under xeric conditions. We also found compelling evidence for strong species‐specific competitive effects in these forests. Altogether, these results constitute critical new information which not only furthers our understanding of important theoretical questions about the assembly of Mediterranean forests, but will also be of help in developing new guidelines for adapting forests in this climatic boundary to global change. If we consider the climatic gradients of this study as a surrogate for future climatic conditions, then we should expect absolute growth rates to decrease and sensitivity to competition to increase in most forests of the Iberian Peninsula (in all but the northern Atlantic forests), making these management considerations even more important in the future.     

Markov models and initial floristic composition in heathland vegetation dynamics

A simple Markov model is used to test the hypothesis that the floristic composition of vegetation colonizing bare ground immediately after burning is the major factor influencing post-fire development in heathland vegetation. Data are taken from stands of different ages at time of burning in a species-rich Calluna-Arctostaphylos heath in NE Scotland. It is shown that variation in the initial floristic composition of the stands is not, in itself, sufficient to produce model simulations which match observed trends, although altering the probabilities of transition from bare ground to other states allows more successful simulations. The model supports the hypothesis that stand age before burning influences the post-fire development through the process of colonization of bare ground. After a very severe fire post-fire development may initially depend on the formation of a moss cover, although this requires further study. It is concluded that simple Markov models can provide the basis for examining successional processes when used in a comparative way.Nomenclature follows Tutin et al. (1964–80) for vascular plants and Smith (1978) for mosses.We thank Prof. C. H. Gimingham for comments on the draft, and J. M. M. Humphrey, P. Marren and the Nature Conservancy Council for permission to work on the Muir of Dinnet.     

Disturbance,competition and the maintenance of clonal diversity in Daphnia pulex

Laboratory microcosm experiments tested the intermediate disturbance hypothesis, which states that the highest level of diversity (e.g. species diversity) will be maintained at intermediate scales of disturbance. The effects of disturbance on the maintenance of clonal diversity and on competitive interactions among clones of the obligately parthenogenetic freshwater cladoceran, Daphnia pulex were examined. No significant effect of disturbance size (i.e. dilution volume) on clonal diversity was noted. However, frequency of disturbance had a pronounced effect on clonal diversity, with the highest clonal diversity maintained at low to intermediate disturbance frequencies. Competitive hierarchies among clones were often invariant within a given experiment. Generally, one or two clones dominated, with several less abundant clones persisting throughout an experiment. Results suggest that low to intermediate disturbances could be important in the maintenance of genetic variation in natural populations (i.e. through pre-emption of competitive exclusion between genotypes). This could have a direct bearing on the maintenance of both intra- and interspecific diversity.     

Disturbance affects short-term facilitation, but not long-term saturation, of exotic plant invasion in New Zealand forest

We investigate the spread of an exotic herb, Hieracium lepidulum, into a New Zealand Nothofagus forest with the aim of understanding how stand-development of tree populations, propagule pressure and invader persistence, affect invasion across the landscape and within communities. Using data repeatedly collected over 35 years, from 250 locations, we parametrize continuous-time Markov chain models and use these models to examine future projections of the invasion under a range of hypothetical scenarios. We found that the probability of invasion into a stand was relatively high following canopy disturbance and that local abundance of Hieracium was promoted by minor disturbances. However, model predictions extrapolated 45 years into the future show that neither the rate of landscape-level invasion, nor local population growth of Hieracium, was affected much by changing the frequency of canopy disturbance events. Instead, invasion levels were strongly affected by the ability of Hieracium to persist in the understorey following forest canopy closure, and by propagule supply from streams, forest edges and plants already established within the stand. Our results show that disturbance frequency has surprisingly little influence on the long-term trajectory of invasion, while invader persistence strongly determines invasion patterns.     

Positive interactions, discontinuous transitions and species coexistence in plant communities

The population and community level consequences of positive interactions between plants remain poorly explored. In this study we incorporate positive resource-mediated interactions in classic resource competition theory and investigate the main consequences for plant population dynamics and species coexistence. We focus on plant communities for which water infiltration rates exhibit positive dependency on plant biomass and where plant responses can be improved by shading, particularly under water limiting conditions. We show that the effects of these two resource-mediated positive interactions are similar and additive. We predict that positive interactions shift the transition points between different species compositions along environmental gradients and that realized niche widths will expand or shrink. Furthermore, continuous transitions between different community compositions can become discontinuous and bistability or tristability can occur. Moreover, increased infiltration rates may give rise to a new potential coexistence mechanism that we call controlled facilitation.     

Strategies for dealing with incomplete information in the modeling of molecular interaction networks

Modelers of molecular interaction networks encounter the paradoxical situation that while large amounts of data are available, these are often insufficient for the formulation and analysis of mathematical models describing the network dynamics. In particular, information on the reaction mechanisms and numerical values of kinetic parameters are usually not available for all but a few well-studied model systems. In this article we review two strategies that have been proposed for dealing with incomplete information in the study of molecular interaction networks: parameter sensitivity analysis and model simplification. These strategies are based on the biologically justified intuition that essential properties of the system dynamics are robust against moderate changes in the value of kinetic parameters or even in the rate laws describing the interactions. Although advanced measurement techniques can be expected to relieve the problem of incomplete information to some extent, the strategies discussed in this article will retain their interest as tools providing an initial characterization of essential properties of the network dynamics.     

On some formulas in a partnership model from the perspective of a semi-Markov process

Many deterministic models of sexually transmitted diseases, as well as population models in general, contain elements of stochastic or statistical reasoning. An example of such a model is that of Dietz and Hadeler (1988) concerning sexually transmitted diseases in which there is partnership formation and dissolution. Among the interesting formulas in this paper, which enter into the analysis of the model, are those for the expected number of partners a male or female has during a lifetime. To a probabilist such formulas suggest the possibility that some stochastic process may be constructed so as to yield these formulas as well as others that may be of interest. The principal purpose of this paper is to demonstrate that such a stochastic process does indeed exist in the form of a three state semi-Markov process in continuous time with stationary laws of evolution and with a one-step density matrix determined by four parameters which were interpreted as constant latent risk functions in the classical theory of competing risks. This construction of a semi-Markov process not only provides a framework for the systematic derivation of the formulas of Dietz and Hadeler but also suggests pathways,for extensions to the age-dependent case.This research was partially supported by NATO Grant D.890350     

Phytoplankton succession and diversity in a warm monomictic,relatively shallow lake: Lake Volvi,Macedonia, Greece

In Lake Volvi, phytoplankton dominance was examined in relation to the main phases of the fluctuating physical state of the water column and nutrient levels. Four stages of algal succession were identified. The successional pattern was, in general, similar from year to year. External disturbances such as increased wind mixing and high floods had the effect of setting the succession back to an earlier stage.Nanoplanktic diatoms and flagellates dominated during the first stages of succession. The succession was running from r-selected species in early stages to K-strategists in summer and towards a mixed community in the terminal stage.Diversity was expressed by the indices of Gleason and Shannon (based on physical units and biomass). Interpretation of species diversity took into account the differential variations of its two components, the number of species and the evenness of their distribution. Diversity was more affected by evenness than by number of species.Diversity increased in late stages of succession corresponding to the complexity of the community (high number of species, high evenness). Physical disturbances influenced positively the diversity. The maximal diversity appeared in the transition periods between the compositional changes generated by disturbances and the true successional development. Disturbances may generate a more uniform distribution of diversity throughout the whole water column.The role of physical disturbances in increasing the phytoplankton diversity may be demonstrated from comparisons of diversity — evenness — species number of the same stages under different meteorological conditions.