Intrinsic inter-specific competition in a guild of tephritid fruit fly parasitoids: Effect of co-evolutionary history on competitive superiority

Tephritid fruit fly parasitoid guilds are dominated by solitary koinobiont species that attack different host stages, but most emerge as adults from host puparia. Previous studies suggest intrinsic competitive superiority by the egg-attacking parasitoid Fopius arisanus (Sonan) against all larval-attacking parasitoids in Hawaii. In this study, we tested the early-acting competitive superiority prediction in relation to the co-evolutionary history of competition between an egg–larval parasitoid (Fopius ceratitivorus Wharton), and each of three larval parasitoids [Psyttalia concolor (Szépligeti), Diachasmimorpha kraussii (Fullaway), and Diachasmimorpha longicaudata (Ashmead)]. F. ceratitivorus and P. concolor share a common origin (eastern Africa), while D. kraussii is an Australian species, and D. longicaudata is from Southeast Asia. The outcomes of intrinsic competition between the egg-attacking parasitoid and each of the three larval-attacking parasitoids within their common host, the Mediterranean fruit fly Ceratitis capitata (Wiedemann) were compared. F. ceratitivorus invariably eliminated the co-evolved P. concolor through physiological suppression of the later-attacking parasitoid’s egg development, providing evidence that supports the early-acting-superiority hypothesis. However, F. ceratitivorus was unable to suppress development of the two non co-evolved larval parasitoids. Instead, the larvae of both later-acting parasitoid species physically killed F. ceratitivorus larvae inside the host. The results suggest that co-evolutionary history influences competitive superiority. The evolution of inter-specific competition and its implications for biological control are discussed.     

Inference from plant competition experiments: the effect of spatial covariance

Two-species plant competition experiments are often drawn on in an attempt to understand and possibly to predict the formation of plant community structures. Frequently, such competition experiments use a non-random spatial design but is analysed in models where a random spatial design is implicitly assumed. Here, the effect of spatial covariance in plant competition experiments on the inferred ecological conclusions is demonstrated. It is shown that spatial covariance, which is dependent on the competition kernel and the experimental design, may affect the estimation of parameters of interest. Based on the same set of experimental data, the predicted outcome of competition between two Arabidopsis thaliana genotypes can vary between (almost certain) coexistence and (almost certain) exclusion of one genotype, depending on the assumptions made about the spatial scale of competition between the plants . The predicted outcome of competition depended strongly on the mean interaction distance of the competition kernel, whereas the functional shape of the competition kernel was less important. It is suggested that plants in manipulated competition experiments should be placed randomly, such that the design of the competition experiment is in agreement with the model used in the analysis.     

Heterotrophic bacterial guild structure: Relationship to biodegradative populations

Numerical taxonomic analysis of a freshwater bacterial guild demonstrated that the bacteria capable of growth on phenanthrene and polychlorinated biphenyl media were representative of the taxa obtained from low nutrient oligotrophic media. The diversity of heterotrophic bacteria and members of new taxa recovered from the guild followed a poisson distribution relative to the number of isolation media used. Moderately high nutrient, yeast extract peptone and glucose agar was found to be the most selective isolation medium relative to the total number of taxa recovered whereas low nutrient, lake water agar was the least selective medium used. Carbon source utilization patterns of the isolated taxa indicated that taxa within the guild had broad niche ranges and could potentially occupy many niches within a dynamic environment. The structure of the bacterial guild was dominated by mesophilic oligotrophs. The results of this investigation demonstrate that potential biodegradative populations are representative of the diverse taxa found in uncontaminated freshwater environments.     

Incompatibility and pollen competition in Alnus glutinosa: Evidence from pollination experiments

Different types of incompatibility systems were found to operate simultaneously in alnus glutinosa in the course of numerous pollination experiments, including self-pollination and pollination with controlled pollen mixtures. Isozyme genetic markers were used to identify the pollen parent of each offspring from the mixed pollination experiments, thus allowing specification of the fertilization success of each pollen parent. In a first step, these results were compared with observations on in vitro pollen germination experiments. This comparison allows for exploration of the explanatory value of different germination media as models of germination conditions on stigmas. In most cases, the data suggest that the in vitro germination conditions resemble the fertilization conditions in vivo, at least in the sense that they favor the same pollen parents. By providing a generic and operable definition of the two basic types of incompatibility, eliminating (inability to fertilize ovules) and cryptic (resulting in lowered fertilization success of a pollen parent under competition), evidence was detected for the existence of both types of incompatibility in alnus glutinosa, where eliminating incompatibility occurred as self-incompatibility only. However, since this incompatibility seems to act primarily via pollen elimination, seed production is not likely to be negatively affected in natural populations, even for comparatively large amounts of self-pollination. This revised version was published online in July 2006 with corrections to the Cover Date.     

Pitfalls in experiments testing predictions from sperm competition theory

As females of many species mate with more than one male, ejaculates often face competition from the sperm of other males. In recent years, numerous papers have been published on theoretical predictions of evolutionary, behavioural and physiological responses to variation in the strength of sperm competition (SC). These theoretical predictions have also been extensively tested. However, although predictions from SC theory are relatively straightforward, extra caution has to be paid in the design of experiments testing them. One difficulty is for example to disentangle immediate and mean SC risk and intensity. Without carefully designed experiments, it is also very easy to simultaneously increase SC risk and the probability of intense SC--a situation for which we currently have no clear predictions, as the theoretical models to date only assume variation in either SC risk or intensity. In this paper, we discuss these and some other pitfalls related to manipulations of SC risk and intensity and suggest how to avoid them.     

Induced resistance in mountain birch: defence against leaf-chewing insect guild and herbivore competition

Summary The effect of leaf damage simulating the feeding of early season insect herbivore species, e.g. Epirrita autumnata, to mountain birch, Betula pubescens ssp. tortuosa, on the performance of insect larvae was studied with eleven leaf-chewing sawfly species. I found variation in the results that was due to short- and long-term inducible responses and to the phenology of herbivore species. In general, early and mid-season species were more strongly affected by induced reactions than late-season species. This finding is in accordance with earlier results but I could show that the persistance of induced reactions rather than the influence of timing of damage is responsible for the result. The growth of the larvae of mid-season sawfly species was affected by both short- and long-term induced reactions. This result shows that early season species may escape short-term induced reactions of mountain birch in current year but may not avoid long-term effects. It is supposed that seasonal deterioration of leaf quality either masks the effects of induced defences or late-season species are better adapted to low-quality leaves. Some species show variation in their response to induced defence in different years. This may be due to yearly differences in induced reactions as well as to species-specific responses. Induced defence reactions may play a role in competitive interactions between herbivore species in leaf-chewing guild of mountain birch.     

Resource partitioning and interspecific competition in snakes: the search for general geographical and guild patterns

The role of interspecific competition as a key factor in the ecology of natural communities where species exploit limited resources is well established, and the study of competition dynamics in snake communities has received much attention in recent years. Twenty years ago, an acclaimed review ( Toft 1985 ) suggested that snakes were atypical among vertebrates because sympatric species usually partition the food niche. Here, I review the articles published in the last two decades with the aim of finding any general geographical or guild patterns and assessing if Toft's main conclusion is still supported by new evidence. Where appropriate, I use Monte Carlo simulations to establish whether observed patterns of niche overlap are real, or if they have occurred by chance. My study shows clear congruence in the patterns of coexistence exhibited by snake communities in different regions of the world, i.e.: (1) cold regions of the northern hemisphere (high latitudes and altitudes) exhibit low species richness and a very low, or even absent, potential for interspecific competition; (2) aquatic snakes that form communities in temperate regions generally partition the food type available and exhibit a broad similarity in habitat use with subtle differences in microhabitat use; (3) terrestrial snake communities in temperate regions are very variable in terms of their coexistence dynamics and show no evidence of generalised patterns; (4) sympatric viperids in Europe, North America and, most interestingly, tropical Asia partition the available habitat but not the prey resource; (5) competition is much stronger in tropical snake communities, and the intensity of this process fluctuates throughout the year being most intense during periods of low food availability; (6) in general, tropical snakes partition the food resource (prey type and/or prey size), but when this resource is not partitioned competitive exclusion can occur. Prey resource availability is a fundamental variable for all snake communities; this is clearly documented by studies on terrestrial snakes in Australia where, due to a relative scarcity of prey availability in the field, sympatry among species is much rarer than in other continents. I conclude that, although there are several notable exceptions, Toft's main conclusion is still supported by empirical evidence. However, I disagree with Toft's conclusion that most snakes are food specialists, and I contend that interspecific competition is important in structuring many (if not most) of the snake communities around the world.     

Statistical methods for estimating complexity from competition experiments between two populations

Often a screening or selection experiment targets a cell or tissue, which presents many possible molecular targets and identifies a correspondingly large number of ligands. We describe a statistical method to extract an estimate of the complexity or richness of the set of molecular targets from competition experiments between distinguishable ligands, including aptamers derived from combinatorial experiments (SELEX or phage display). In simulations, the non-parametric statistic provides a robust estimate of complexity from a 100 ×100 matrix of competition experiments, which is clearly feasible in high-throughput format. The statistic and method are potentially applicable to other ligand binding situations.     

Replacement interaction diagrams in interspecific competition experiments

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

Estimating relative fitness in viral competition experiments

The relative fitness of viral variants has previously been defined as the slope of the logarithmic ratio of the genotype or phenotype frequencies in time plots of pairwise competition experiments. Developing mathematical models for such experiments by employing the conventional coefficient of selection s, we demonstrate that this logarithmic ratio gives the fitness difference, rather than the relative fitness. This fitness difference remains proportional to the actual replication rate realized in the particular experimental setup and hence cannot be extrapolated to other situations. Conversely, the conventional relative fitness (1 + s) should be more generic. We develop an approach to compute the generic relative fitness in conventional competition experiments. This involves an estimation of the total viral replication during the experiment and requires an estimate of the average lifetime of productively infected cells. The novel approach is illustrated by estimating the relative fitness, i.e., the relative replication rate, of a set of zidovudine-resistant human immunodeficiency virus type 1 variants. A tool for calculating the relative fitness from observed changes in viral load and genotype (or phenotype) frequencies is publically available on the website at http://www-binf.bio.uu.nl/( approximately )rdb/fitness.html.     

Application of FAME (fatty acid methyl ester) analysis in the numerical taxonomic determination of bacterial guild structure

Comparative numerical taxonomic analyses, using fatty acid methyl ester (FAME) profiles and phenetic characteristics, were conducted to examine bacterial guild structure in freshwater sediments. Both approaches were used to examine a subset of 60 OTUs obtained from a previously well characterized microbial community in sediments of a shallow fast flowing stream. For both classifications, greater than 80% of the OTUs were recovered in 11 and 12 major groups for FAME and phenetics approaches. However, there was not complete correspondence for the groupings of the 2 classifications, with most FAME groups being distributed among the phenetic groups and 2 phenetic groups not providing usable characteristics for FAME analysis. The results did demonstrate significant taxonomic variation in bacteria capable of occupying the same or a similar fundamental niche.     

Interspecific relations in forest floor coleopteron assemblages: Niche overlap and guild structure

In six different environments of the Towada-Hachimantai National Park, the forest floor coleopterons were sampled by means of pitfall traps with five different baits. Their niche breadths and niche overlaps were assessed for the electivity of habitat, bait, and habitat × bait, and compared with those expected from neutral models constructed randomly. The niche breadths in the observed assemblages were smaller than those expected from the neutral model, while the niche overlaps were more widely distributed than those expected from the neutral model. Mean overlaps between species neighbors in niche space, cluster analyses, and ordinations of species revealed a guild for habitat, bait and habitat × bait. Lawlor's (1979) gamma matrices concerning the habitat electivity show that the rates of mutualism resulted from indirect interspecific competitions are significantly smaller in the guild than in the whole assemblage. This suggests that the interspecific competition among the guild members is mitigated or even converted into mutualism by joining of the species which do not belong to the guild. The rates of mutualism were also significantly larger in the resource state of two dimensions (habitat × bait) than in that of either one. Therefore, if niche overlaps are measured on the basis of resource state of multi-dimensions, the rates of mutualism may increase further.     

The mechanisms of interference competition: two experiments on foraging waders

Models of population dynamics that include interference competitionhave often been applied to foraging waders and less so to otherforagers, even though these models are, in principle, generallyapplicable. At present, however, it is still unclear whetherinterference competition is of importance for foraging waders.To support this idea experimental evidence and knowledge ofthe mechanisms underlying interference effects are required.We experimentally determined the relationship between foragerdensity and foraging success in two wader species: the red knot(Calidris canutus) and the ruddy turnstone (Arenaria interpres).With each of the two species, we conducted an experiment consistingof 300 one-min trials. In these trials we scored the behaviorand the foraging success of focal individuals at specific combinationsof bird and prey density. Irrespective of prey density, individualsof both species discovered fewer prey items at higher bird densities.Despite this, only in turnstones did intake rates decline withincreasing bird density. Knots compensated for a lower prey-discoveryrate by rejecting fewer prey items at higher bird densities.In knots, bird density had a complex, nonmonotonic effect onthe time spent vigilant and searching. In turnstones the maineffect of increased bird density was a reduction in the prey-encounterrate, that is, the reward per unit search time. Effects on thetime spent vigilant and the time spent searching were less pronouncedthan in knots. Thus, the mechanistic basis of the effects ofbird density was complex for each of the two species and differedbetween them.     

Experimental determination of predation intensity in an intertidal predator guild: dominant versus subordinate prey

Theoretical and empirical ecologists have long acknowledged that information about the intensity or strength of the interaction between species is crucial for an understanding of community dynamics. In communities in which predation is an important structuring process, and some predator species are commercially exploited, quantitative estimates of predation by different predator species within a guild are necessary to make even the simplest recommendations about conservation and resource management. Here, we evaluated per capita and population level components of predation intensity of three intertidal predators that feed on monospecific stands of barnacles and mussels at wave exposed sites in the rocky intertidal zone of central Chile. These prey species represent the two most distinctive stages of the mid-intertidal seascape, with mussels being competitively dominant. Our results showed that the commercially exploited gastropod Concholepas concholepas and the sea star Heliaster helianthus have similarly large per capita and population effects on the competitively dominant mussel Perumytilus purpuratus . Their per capita (by average size individual) and population effects on mussels were more than two orders of magnitude larger than those of Acanthocyclus gayi crabs and likely even larger than the effect of other predator species in this system (other crabs, whelks, birds, fish). The overall pattern of predation on barnacles was similar to that on mussels, but some differences occurred in the way different components of predation intensity were distributed across predator species. Despite the roughly similar pattern of population predation intensity between prey species, the expected consequences for the prey population, and hence the rest of the community, were acutely different for mussels and barnacles.     

Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

Summary A particular linear programming model is constructed to predict the diets of each of 14 species of generalist herbivores at the National Bison Range, Montana. The herbivores have body masses ranging over seven orders of magnitude and belonging to two major taxa: insects and mammals. The linear programming model has three feeding constraints: digestive capacity, feeding time and energy requirements. A foraging strategy that maximizes daily energy intake agrees very well with the observed diets. Body size appears to be an underlying determinant of the foraging parameters leading to diet selection. Species that possess digestive capacity and feeding time constraints which approach each other in magnitude have the most generalized diets. The degree that the linear programming models change their diet predictions with a given percent change in parameter values (sensitivity) may reflect the observed ability of the species to vary their diets. In particular, the species which show the most diet variability are those whose diets tend to be balanced between monocots and dicots. The community-ecological parameters of herbivore body-size ranges and species number can possibly be related to foraging behavior.     

Automated structure determination from NMR spectra

Automated methods for protein structure determination by NMR have increasingly gained acceptance and are now widely used for the automated assignment of distance restraints and the calculation of three-dimensional structures. This review gives an overview of the techniques for automated protein structure analysis by NMR, including both NOE-based approaches and methods relying on other experimental data such as residual dipolar couplings and chemical shifts, and presents the FLYA algorithm for the fully automated NMR structure determination of proteins that is suitable to substitute all manual spectra analysis and thus overcomes a major efficiency limitation of the NMR method for protein structure determination.     

Variability in species richness and guild structure in two species-rich grasslands

It has been suggested that variation in the proportion of species in guilds (=guild proportionality) indicates community structuring by guilds in biotic communities. This hypothesis was tested on a subthermophilous grassland and a mesotrophic meadow at a scale of 0.09 m² based on a five-year data set. Further, variation in the total number of species, variation in the number of species belonging to a guild and non-randomness in species composition of guilds were studied. A number of criteria for guild definition were used, such as life form, Grime's C-S-R strategy, phenology, plant height, pollination and dispersal syndromes, leaf shape and anatomy and taxonomy at the family level. The observed variation in the number of guild species corresponded to the null model in which species assemblages with fixed species richness per square were randomly generated from the species pool. The observed variation in the number of guild species was often higher than the variation calculated for randomly distributed species whereas the variation in the proportion of guild species was in some cases lower than the variation calculated for randomly distributed species with fixed frequencies. Possible reasons for the discrepancy in the results based on different models are discussed. It is concluded that there is little evidence of guilds in the organization of grasslands. *** DIRECT SUPPORT *** A02DO006 00012     

Niches, rather than neutrality, structure a grassland pioneer guild

Pioneer species are fast-growing, short-lived gap exploiters. They are prime candidates for neutral dynamics because they contain ecologically similar species whose low adult density is likely to cause widespread recruitment limitation, which slows competitive dynamics. However, many pioneer guilds appear to be differentiated according to seed size. In this paper, we compare predictions from a neutral model of community structure with three niche-based models in which trade-offs involving seed size form the basis of niche differentiation. We test these predictions using sowing experiments with a guild of seven pioneer species from chalk grassland. We find strong evidence for niche structure based on seed size: specifically large-seeded species produce fewer seeds but have a greater chance of establishing on a per-seed basis. Their advantage in establishment arises because there are more microsites suitable for their germination and early establishment and not directly through competition with other seedlings. In fact, seedling densities of all species were equally suppressed by the addition of competitors' seeds. By the adult stage, despite using very high sowing densities, there were no detectable effects of interspecific competition on any species. The lack of interspecific effects indicates that niche differentiation, rather than neutrality, prevails.