Resource-Ratio Theory and the Control of Invasive Plants

There have been relatively few tests of resource-ratio theory in terrestrial systems. Additionally, resources are known to be an important factor determining the success of invasive species. Here I discuss how the study by Newingham and Belnap (pp. 29–40, this issue) tests predictions of resource-ratio theory and how they apply it to questions of invasion by Bromus tectorum in a terrestrial grassland.     

Plant competition and exclusion with optimizing individuals

Most models of plant competition represent competition as taking place between species when realistically competition takes place between individuals. We model individual plants as optimally choosing biomass in order to maximize net energy that is directed into reproduction. Competition is for access to light and a plant that grows more biomass adds to the leaf area index, creating negative feedback in the form of more self shading and shading of its neighbors. In each period and for given species densities, simultaneous maximization by all plants yields an equilibrium characterized by optimum biomasses. Between periods the net energies plants obtain are used to update the densities, and if densities change the equilibrium changes in the subsequent period. A steady state is attained when all plants have net energies that just allow for replacement. Four main predictions of the resource-ratio theory of competition are obtained, providing behavioral underpinnings for species level models. However, if individual plant parameters are not identical across species, then the predictions do not follow. The optimization framework yields many other predictions, including how specific leaf areas and resource stress impact biomass and leaf area indices.     

Resource heterogeneity, soil fertility, and species diversity: effects of clonal species on plant communities

Spatial heterogeneity in soil resources is widely thought to promote plant species coexistence, and this mechanism figures prominently in resource-ratio models of competition. However, most experimental studies have found that nutrient enhancements depress diversity regardless of whether nutrients are uniformly or heterogeneously applied. This mismatch between theory and empirical pattern is potentially due to an interaction between plant size and the scale of resource heterogeneity. Clonal plants that spread vegetatively via rhizomes or stolons can grow large and may integrate across resource patches, thus reducing the positive effect of small-scale resource heterogeneity on plant species richness. Many rhizomatous clonal species respond strongly to increased soil fertility, and they have been hypothesized to drive the descending arm of the hump-shaped productivity-diversity relationship in grasslands. We tested whether clonals reduce species richness in a grassland community by manipulating nutrient heterogeneity, soil fertility, and the presence of rhizomatous clonal species in a 6-year field experiment. We found strong and consistent negative effects of clonals on species richness. These effects were greatest at high fertility and when soil resources were applied at a scale at which rhizomatous clonals could integrate across resource patches. Thus, we find support for the hypothesis that plant size and resource heterogeneity interact to determine species diversity.     

Theoretical predictions for how temperature affects the dynamics of interacting herbivores and plants

Concern about climate change has spurred experimental tests of how warming affects species' abundance and performance. As this body of research grows, interpretation and extrapolation to other species and systems have been limited by a lack of theory. To address the need for theory for how warming affects species interactions, we used consumer-prey models and the metabolic theory of ecology to develop quantitative predictions for how systematic differences between the temperature dependence of heterotrophic and autotrophic population growth lead to temperature-dependent herbivory. We found that herbivore and plant abundances change with temperature in proportion to the ratio of autotrophic to heterotrophic metabolic temperature dependences. This result is consistent across five different formulations of consumer-prey models and over varying resource supply rates. Two models predict that temperature-dependent herbivory causes primary producer abundance to be independent of temperature. This finding contradicts simpler extensions of metabolic theory to abundance that ignore trophic interactions, and is consistent with patterns in terrestrial ecosystems. When applied to experimental data, the model explained 77% and 66% of the variation in phytoplankton and zooplankton abundances, respectively. We suggest that metabolic theory provides a foundation for understanding the effects of temperature change on multitrophic ecological communities.     

Stoichiometric constraints on resource use, competitive interactions, and elemental cycling in microbial decomposers

Heterotrophic microbial decomposers, such as bacteria and fungi, immobilize or mineralize inorganic elements, depending on their elemental composition and that of their organic resource. This fact has major implications for their interactions with other consumers of inorganic elements. We combine the stoichiometric and resource-ratio approaches in a model describing the use by decomposers of an organic and an inorganic resource containing the same essential element, to study its consequences on decomposer interactions and their role in elemental cycling. Our model considers the elemental composition of organic matter and the principle of its homeostasis explicitly. New predictions emerge, in particular, (1) stoichiometric constraints generate a trade-off between the R* values of decomposers for the two resources; (2) they create favorable conditions for the coexistence of decomposers limited by different resources and with different elemental demands; (3) however, combined with conditions on species-specific equilibrium limitation, they draw decomposers toward colimitation by the organic and inorganic resources on an evolutionary time scale. Moreover, we derive the conditions under which decomposers switch from consumption to excretion of the inorganic resource. We expect our predictions to be useful in explaining the community structure of decomposers and their interactions with other consumers of inorganic resources, particularly primary producers.     

Interactive effects of productivity and predation on zooplankton diversity

Recent studies suggest the necessity of understanding the interactive effects of predation and productivity on species coexistence and prey diversity. Models predict that coexistence of prey species with different competitive abilities can be achieved if inferior resource competitors are less susceptible to predation and if productivity and/or predation pressure are at intermediate levels. Hence, predator effects on prey diversity are predicted to be highly context dependent: enhancing diversity from low to intermediate levels of productivity or predation and reducing diversity of prey at high levels of productivity or predation. While several studies have examined the interactive effects of herbivory and productivity on primary producer diversity, experimental studies of such effects in predator‐prey systems are rare. We tested these predictions using an aquatic field mesocosm experiment in which initial density of the zooplankton predator Notonecta undulata and productivity were manipulated to test their interactive effects on diversity of seven zooplankton, cladoceran species that were common in surrounding ponds. Two productivity levels were imposed via phosphorus enrichment at levels comparable to low and intermediate levels found within neighboring natural ponds. We used open systems to allow for natural dispersal and behaviorally‐mediated numerical responses by the flight‐capable predator. Effects of predators on zooplankton diversity depended on productivity level. At low and high productivity, prey species richness declined while at high productivity it showed a unimodal relationship with increasing the predator density. Effects of treatments were weaker when using Pielou's evenness index or the inverse Simpson index as measures of prey diversity. Our findings are generally consistent with model predictions in which predators can facilitate prey coexistence and diversity at intermediate levels of productivity and predation intensity. Our work also shows that the functional form of the relationship between prey diversity and predation intensity can be complex and highly dependent on environmental context.     

A test of the niche dimension hypothesis in an arid annual grassland

The niche dimension hypothesis predicts that greater numbers of limiting factors can allow greater numbers of species to coexist through species' tradeoffs for different limiting factors. A prediction that follows is that addition of multiple limiting resources to plant communities will increase productivity and simultaneously decrease diversity. Species loss due to limiting resource enrichment might occur through reducing the number of resources that species compete for or by changing the identity of limiting factors. We tested these predictions of the niche dimension hypothesis in an arid annual grassland by adding combinations of nutrients: nitrogen (N), phosphorus (P), and potassium with other elements (O). We found that species number decreased while biomass increased with greater numbers of added resources. In particular, N in combinations with P or O resulted in the greatest species loss, while biomass increased super-additively with N and P together. The addition of greater numbers of added nutrients decreased the availability of light and soil moisture, consistent with a potential shift in the identity of limiting resources. Species also differed in their responses to different combinations of N, P, and O, supporting predictions of resource-ratio tradeoffs. These results are particularly notable because this experiment was conducted during a drought year in an arid grassland (226 mm annual rainfall), which might have been expected to be water-rather than nutrient-limited. Our results support the hypothesis that plant diversity may be maintained by high-dimensional tradeoffs among species in their abilities to compete for multiple limiting factors.     

Dispersal and diversity in experimental metacommunities: linking theory and practice

There has been a recent rise in the number of experiments investigating the effect of dispersal on diversity, with many of the predictions for these tests derived from metacommunity theory. Despite the promise of linking observed relationships between dispersal and diversity to underlying metacommunity processes, empirical studies have faced challenges in providing robust tests of theory. We review experimental studies that have tested how dispersal affects metacommunity diversity to determine why shortcomings emerge, and to provide a framework for empirical tests of theory that capture the processes structuring diversity in natural metacommunities. We first summarize recent experimental work to outline trends in results and to highlight common methods that cause a misalignment between empirical studies and the processes described by theory. We then identify the undesired implications of three widely used experimental methods that homogenize metacommunity structure or species traits, and present alternative methods that have been used to successfully integrate experiments and theory in a biologically relevant way. Finally, we present methodological and theoretical insights from three related ecological fields (coexistence, food web and priority effects theory) that, if integrated into metacommunity experiments, could help isolate the independent and joint effects of local interactions and dispersal on diversity, and reveal the mechanisms underlying observed dispersal–diversity patterns. Together, these methods can provide stronger tests of existing theory and stimulate new theoretical explorations. Synthesis Although metacommunity experiments offer a unique opportunity to test classic and emerging theory on the relationship between dispersal and diversity, several common challenges have hindered robust tests of theory. We outline how emerging theory on the invasion criterion, food webs and priority effects could be help clarify when and how dispersal affects metacommunity diversity, and identify when experimental approaches that homogenize metacommunities fail to test existing theory. By forging better links between theoretical and empirical work, we hope to motivate novel and improved experimental approaches to understanding the joint effects of local and regional processes on diversity.     

资源竞争理论及其研究进展

介绍了国外近20年来发展起来的一种新的竞争理论——资源竞争理论。该理论包含两个主要假说,即R^*-法则和资源比假说。资源竞争理论已在微生物、微藻、高等植物及浮游动物中得到广泛的实验验证。阐述了资源竞争理论形成的基础——Monod竞争模型和Droop竞争模型。对两种模型在稳态及非稳态条件下的预测性能作了比较：对于稳态下的竞争,两种模型皆能做出较好的预测,但使用Monod模型更为简便;对于非稳态下的竞争,使用Droop模型更为合理。对资源竞争理论的发展趋势作了展望。     

Density-dependent habitat selection: evaluating isoleg theory with a Lotka-Volterra model

Contemporary models of density-dependent habitat selection generally focus on long-term evolutionary consequences of intraspecific or interspecific competition and/or patterns of resource use in patchy environments. A primary goal of such studies often is to elucidate evolutionary stable strategies based on steady-state dynamics of population growth. In contrast, we developed a simulation model to explore short-term movements of interspecific competitors among fine-grained habitats of differing attributes, as might result from field manipulations of habitat quality or population densities. In this model, habitat quality is expressed in terms of mean individual fitness, represented by average per capita growth rate calculated according to the Lotka-Volterra equations describing interspecific competition. This model provides a mechanism for quantifying the effects of habitat quality, patterns of resource use and competition on distributions of individuals. Results demonstrate the heuristic value of this model in corroborating predictions derived from the ideal free distribution and isodar theory, and in generating isolegs to test the predictions of isoleg theory. Results indicate that small changes in model parameters have substantial impacts on patterns of habitat use and co-occurrence between species. The model identifies a variety of conditions under which isolegs for a given type of community organization deviate from predictions of contemporary isoleg theory, potentially expanding the universe of possible interspecific behaviors underlying the development of evolutionary stable strategies.     

Non-neutral community dynamics: empirical predictions for ecosystem function and diversity from linearized consumer–resource interactions

A general model of linearized species interactions, essentially Lotka–Volterra theory, applied to questions of biodiversity has previously been shown to be a powerful tool for understanding local species–abundance patterns and community responses to environmental change for a single trophic level. Here this approach is extended to predict community composition and responses to environmental changes in trophically structured systems. We show how resource and consumer species richness and their relative abundances vary with the means and variances in enrichment level and strengths of intra- and interspecific interactions. Also demonstrated are the responses of local resource and consumer species richness to the global species pools at both trophic levels, as well as the covariation with net resource productivity. These predictions for resource and consumer specific responses to changes in environmental enrichment and global biodiversity are directly testable.     

Social structure and facultative mating systems of hoary marmots (Marmota caligata)

Mate-choice theory predicts different optimal mating systems depending on resource availability and habitat stability. Regions with limited resources are thought to promote monogamy. We tested predictions of monogamy in a social rodent, the hoary marmot (Marmota caligata), at the northern climatic extreme of its distribution. Mating systems, social structure and genetic relationships were investigated within and among neighbouring colonies of marmots within a 4 km(2) valley near Kluane National Park, Yukon, Canada, using 21 microsatellite loci. While both monogamous and polygynous populations of hoary marmots have been observed in the southern reaches of this species' range; northern populations of this species are thought to be predominantly monogamous. Contrary to previous studies, we did not find northern hoary marmot social groups to be predominantly monogamous; rather, the mating system seemed to be facultative, varying between monogamy and polygyny within, as well as among, social groups. These findings reveal that the mating systems within colonies of this species are more flexible than previously thought, potentially reflecting local variation in resource availability.     

A dynamic model of facilitation on environmental stress gradients

Theories based on competition for resources in animals and other non‐sessile organisms rarely consider the role of facilitative interactions. Yet these interactions are important for community assembly, especially under stressful environments (e.g. the stress‐gradient hypothesis, SGH). To make an explicit link between species interaction theory and SGH patterns, I used a classic resource competition model promoting coexistence between a beneficiary and its facilitator sharing a common resource along a stress gradient. I compared model outcomes for two fundamentally different mechanisms of facilitation (alleviation of resource versus non‐resource stress), and also tested the effect of a reciprocal cost of facilitation from the beneficiary. I then tested model's biological relevance using experimental data from two tuber moth species (Lepidoptera, Gelechiidae) for which facilitation in resource access was previously established. Simulation outcomes revealed that both the mode of facilitation and the incorporation of facilitation costs affected the shape of the facilitation–stress relationship. These predictions are in line with current SGH observations and experiments on both plants and animals and reconcile the frequently reported variability of this relationship in nature. Moreover, a sensitivity analysis of model's parameters confirmed the robustness of the modelling framework to uncover the mechanisms responsible for observed species interaction–stress patterns. Finally, when parameterized with tuber moth demographic data, model's results corresponded to observed interaction outcomes along resource stress gradients. Overall, having a common model for plants and animals may simplify assumptions in SGH studies, allow contrasting the shapes of different consumer–resource relationships and specifying the conditions that favour one type of interaction outcome over another.     

Putting competition strategies into ideal free distribution models: habitat selection as a tug of war

When resources are patchily distributed in an environment, behavioral ecologists frequently turn to ideal free distribution (IFD) models to predict the spatial distribution of organisms. In these models, predictions about distributions depend upon two key factors: the quality of habitat patches and the nature of competition between consumers. Surprisingly, however, no IFD models have explored the possibility that consumers modulate their competitive efforts in an evolutionarily stable manner. Instead, previous models assume that resource acquisition ability and competition are fixed within species or within phenotypes. We explored the consequences of adaptive modulation of competitive effort by incorporating tug-of-war theory into payoff equations from the two main classes of IFD models (continuous input (CI) and interference). In the models we develop, individuals can increase their share of the resources available in a patch, but do so at the costs of increased resource expenditures and increased negative interactions with conspecifics. We show how such models can provide new hypotheses to explain what are thought to be deviations from IFDs (e.g., the frequent observation of fewer animals than predicted in "good" patches of habitat). We also detail straightforward predictions made uniquely by the models we develop, and we outline experimental tests that will distinguish among alternatives.     

Two gerbils of the Negev: A long-term investigation of optimal habitat selection and its consequences

Optimal foraging theory has entered a new phase. It is not so much tested as used. It helps behavioural ecologists discover the nature of the information in an animals brain. It helps population ecologists reveal coefficients of interaction and their patterns of density-dependent variation. And it helps community ecologists examine niche relationships. In our studies on two species of Negev desert gerbil, we have taken advantage of the second and third of these functions. Both these gerbils prefer semi-stabilized dune habitat, and both altered their selective use of this habitat and stabilized sand according to experimental changes we made in their populations. Their changes in selectivity agree with a type of optimal foraging theory called isoleg theory. Isoleg theories provide examples of dipswitch theories – bundles of articulated qualitative predictions – that are easier to falsify than single qualitative predictions. By linking behaviour to population dynamics through isoleg theory, we were able to use the behaviour of the gerbils to reveal the shapes of their competitive isoclines. These have the peculiar non-linear shapes predicted by optimal foraging theory. Finally, when owl predation threatens, the behaviour of Gerbillus allenbyi reveals the shape of their victim isocline. As has long been predicted by predation theory and laboratory experiments, it is unimodal.     

Limited versus unlimited membership in microbial communities: evaluation and experimental tests of some paradigms

Recent developments in community ecology have allowed for the synthesis of community models based on principles of limited and unlimited membership. In this discussion, these developments are used as a framework for evaluating the validity of three paradigms that have constrained research on aquatic microbial communities. Because microbes are considered to possess global distributions, species availability is not generally considered to be an important factor determining microbial community composition in most habitats. Requirements for the global distribution of a species are not the same as those for unlimited availability. Rates of propagule transport to isolated and newly formed aquatic systems ( 4 years old) are low enough to have a strong effect on microbial community composition. Natural aquatic systems may require several years to accumulate a full complement of species adapted to environmental conditions at a particular time. Except under extreme circumstances, environmental conditions are not considered to constrain membership in aquatic microbial communities. Most evidence for this contention is based on an inability to detect simple relationships between species distributions and levels of individual environmental parameters. Environmental measurements are often made at a spatial scale much greater than that of the local environment of microbes. Biotic interactions, such as competition, are generally considered to be the predominant force structuring aquatic microbial communities. Although there is an extensive laboratory database to suggest the importance of different types of species interactions, there have been few field studies to confirm this. A general research protocol is described to test predictions derived from current theory of microbial community organization. A mesocosm approach is advocated in order to incorporate crucial aspects of environmental realism into experimental designs while maintaining some of the control found in the laboratory.     

Dos and don'ts of testing the geographic mosaic theory of coevolution

The geographic mosaic theory of coevolution is stimulating much new research on interspecific interactions. We provide a guide to the fundamental components of the theory, its processes and main predictions. Our primary objectives are to clarify misconceptions regarding the geographic mosaic theory of coevolution and to describe how empiricists can test the theory rigorously. In particular, we explain why confirming the three main predicted empirical patterns (spatial variation in traits mediating interactions among species, trait mismatching among interacting species and few species-level coevolved traits) does not provide unequivocal support for the theory. We suggest that strong empirical tests of the geographic mosaic theory of coevolution should focus on its underlying processes: coevolutionary hot and cold spots, selection mosaics and trait remixing. We describe these processes and discuss potential ways each can be tested.     

Species–area relationships across four trophic levels – decreasing island size truncates food chains

That larger areas will typically host more diverse ecological assemblages than small ones has been regarded as one of the few fundamental ‘laws’ in ecology. Yet, area may affect not only species diversity, but also the trophic structure of the local ecological assemblage. In this context, recent theory on trophic island biogeography offers two clear‐cut predictions: that the slope of the species–area relationship should increase with trophic rank, and that food chain length (i.e. the number of trophic levels) should increase with area. These predictions have rarely been verified in terrestrial systems. To offer a stringent test of key theory, we focused on local food chains consisting of trophic specialists: plants, lepidopteran herbivores, and their primary and secondary parasitoids. For each of these four trophic levels, we surveyed species richness across a set of 20 off‐shore continental islands spanning a hundred‐fold range in size. We then tested three specific hypotheses: that species richness is affected by island size, that the slope of the species–area curve is related to trophic rank, and that such differences in slope translate into variation in food chain length with island size. Consistent with these predictions, estimates of the species–area slope steepened from plants through herbivores and primary parasitoids to secondary parasitoids. As a result of the elevated sensitivity of top consumers to island size, food chain length decreased from large to small islands. Since island size did not detectably affect the ratio between generalists and specialists among either herbivores (polyphages vs oligophages) or parasitoids (idiobionts vs koinobionts), the patterns observed seemed more reflective of changes in the overall number of nodes and levels in local food webs than of changes in their linking structure. Overall, our results support the trophic‐level hypothesis of island biogeography. Per extension, they suggest that landscape modification may imperil food web integrity and vital biotic interactions.     

Sex ratio strategies and the evolution of cue use

Quantitative tests of sex allocation theory have often indicated that organism strategies deviate from model predictions. In pollinating fig wasps, Lipporrhopalum tentacularis, whole fig (brood) sex ratios are generally more female-biased than predicted by local mate competition (LMC) theory where females (foundresses) use density as a cue to assess potential LMC. We use microsatellite markers to investigate foundress sex ratios in L. tentacularis and show that they actually use their clutch size as a cue, with strategies closely approximating the predictions of a new model we develop of these conditions. We then provide evidence that the use of clutch size as a cue is common among species experiencing LMC, and given the other predictions of our model argue that this is because their ecologies mean it provides sufficiently accurate information about potential LMC that the use of other more costly cues has not evolved. We further argue that the use of these more costly cues by other species is due to the effect that ecological differences have on cue accuracy. This implies that deviations from earlier theoretical predictions often indicate that the cues used to assess environmental conditions differ from those assumed by models, rather than limits on the ability of natural selection to produce "perfect" organisms.     

The roles of ecological and evolutionary influences in providing structure to parasite species assemblages.

Parasite species assemblages currently are thought to range from isolationist to interactive, their dynamic properties being related to the number of species and types of hosts involved. The literature contains few experimental tests of this concept, however, and many of the host/parasite systems studied to date are not amenable to experimental manipulation. In this review, the presence of a parasite species, in a sample of host individuals, is considered to be an evolutionary phenomenon, but the parasite's population structure is considered to be an ecological one. Studies that allow evaluation of these 2 influences are comparative in nature and include data from a series of homogeneous samples of host populations. A lottery model is presented, in which hosts acquire their assemblages of parasites by Monte Carlo type sampling from multiple kind arrays; the major structuring influence is the relative probability of becoming infected by various parasite species. Claims of parasite species interaction need to be supported by studies showing departures from the predictions of this model. The species density and infraassemblage diversity index distributions are recommended as quantitative tools useful in such work.