Community assembly and diversification in Indo-Pacific coral reef fishes

Theories of species coexistence have played a central role in ecology and evolutionary studies of the origin and maintenance of biodiversity in highly diverse communities. The concept of niche and associated theories predict that competition for available ecological space leads to a ceiling in species richness that influences further diversification patterns. By contrast, the neutral theory supports that speciation is stochastic and diversity independent. We examined the phylogenetic community structure and diversification rates in three families and 14 sites within coral reef fish communities from the Indian and Pacific oceans. Using the phylogenetic relationships among 157 species estimated with 2300 bp of mitochondrial DNA, we tested predictions in terms of species coexistence from the neutral and niche theories. At the regional scale, our findings suggest that phylogenetic community structure shifts during community assembly to a pattern of dispersion as a consequence of allopatric speciation in recent times but overall, variations in diversification rates did not relate with sea level changes. At the local scale, the phylogenetic community structure is consistent with a neutral model of community assembly since no departure from a random sorting of species was observed. The present results support a neutral model of community assembly as a consequence of the stochastic and unpredictable nature of coral reefs favoring generalist and sedentary species competing for living space rather than trophic resources. As a consequence, the observed decrease in diversification rates may be seen as the result of a limited supply of living space as expected in a finite island model.     

Cheating and the evolutionary stability of mutualisms

Interspecific mutualisms have been playing a central role in the functioning of all ecosystems since the early history of life. Yet the theory of coevolution of mutualists is virtually nonexistent, by contrast with well-developed coevolutionary theories of competition, predator-prey and host-parasite interactions. This has prevented resolution of a basic puzzle posed by mutualisms: their persistence in spite of apparent evolutionary instability. The selective advantage of 'cheating', that is, reaping mutualistic benefits while providing fewer commodities to the partner species, is commonly believed to erode a mutualistic interaction, leading to its dissolution or reciprocal extinction. However, recent empirical findings indicate that stable associations of mutualists and cheaters have existed over long evolutionary periods. Here, we show that asymmetrical competition within species for the commodities offered by mutualistic partners provides a simple and testable ecological mechanism that can account for the long-term persistence of mutualisms. Cheating, in effect, establishes a background against which better mutualists can display any competitive superiority. This can lead to the coexistence and divergence of mutualist and cheater phenotypes, as well as to the coexistence of ecologically similar, but unrelated mutualists and cheaters.     

Species coexistence, intransitivity, and topological variation in competitive tournaments

Competitive intransitivity occurs when species’ competitive abilities cannot be listed in a strict hierarchy, but rather form competitive loops, as in the game ‘Rock-Paper-Scissors’. Indices are useful for summarizing intransitivity in communities; however, as with most indices, a great deal of information is compressed into single number. So while recent ecological theory, experiments, and natural history observations demonstrate that competitive intransitivity can promote species coexistence, the consequence of variation in the ‘topology’ of competitive interactions that is not accounted for by intransitivity indices is much less well understood. We use a continuous analytical model and two complementary discrete lattice models (one spatially explicit, the other aspatial) to demonstrate that such variation does indeed greatly affect species coexistence. Specifically, we show that although intransitivity indices are good at capturing broad patterns of coexistence, communities with different levels of intransitivity can have equal coexistence, and communities with equal intransitivity can have different coexistence, due to underlying variation in competitive network topology.     

The importance of niche differentiation for coexistence on large scales

It is widely accepted that niche differentiation plays a key role in coexistence on relatively small scales. With regard to a large community scale, the recently propounded neutral theory suggests that species abundances are more influenced by history and chance than they are by interspecies competition. This inference is mainly based on the probability that competitive exclusion is largely slowed by recruitment limitation, which may be common in species rich communities. In this respect, a theoretical study conducted by Hurtt and Pacala (1995) for a niche differentiated community has been frequently cited to support neutral coexistence. In this paper, we focused on the effect of symmetric recruitment limitation on delaying species competitive exclusion caused by both symmetric and asymmetric competition in a large homogeneous habitat. By removing niche differentiation in space, we found that recruitment limitation could delay competitive exclusion to some extent, but the effect was rather limited compared to that predicted by Hurtt and Pacala's model for a niche differentiated community. Our results imply that niche differentiation may be important for species coexistence even on large scales and this has already been confirmed in some species rich communities.     

Emerging niche clustering results from both competition and predation

Understanding species coexistence has been a central question in ecology for decades, and the notion that competing species need to differ in their ecological niche for stable coexistence has dominated. Recent theoretical and empirical work suggests differently. Species can also escape competitive exclusion by being similar, leading to clusters of species with similar traits. This theory has so far only been explored under competition. By combining mathematical and numerical analyses, we reveal that competition and predation are equally capable to promote clusters of similar species in prey–predator communities, their relative importance being modulated by resource availability. We further show that predation has a stabilizing effect on clustering patterns, making the clusters more diverse. Our results merge different ecological theories and bring new light to the emergent neutrality theory by adding the perspective of trophic interactions. These results open new perspectives to the study of trait distributions in ecological interaction networks.     

Parasites of mutualisms

Cooperation invites cheating, and nowhere is this more apparent than when different species cooperate, known as mutualism. In almost all mutualisms studied, specialist parasites have been identified that purloin the benefits that one mutualist provides another. Explaining how parasites are kept from driving mutualisms extinct remains an unsolved problem because existing theories explaining the maintenance of cooperation do not apply to parasites of mutualisms. Nonetheless, these theories can be summarized in such a way as to suggest how mutualisms can persist in the face of parasites. (1) For cooperation to occur, the recipient of a benefit must reciprocate, and the recriprocated benefit must be captured by the initial giver or its offspring. (2) For cooperation to persist, the mutualism must be re-assembled each generation. Because most mutualisms are of the "by-product' type, broadly defined, the first condition is normally always fulfilled. Thus, the maintenance of mutualism usually requires enforcement of the second condition: reliable re-assembly. Hence, I argue that the persistence of mutualism is best understood by using theories of species coexistence, because each mutualist can be considered a resource for the other, and species coexistence theory explains how multiple taxa (e.g. parasites and mutualists) can stably partition a resource over multiple generations. This approach connects the study of mutualism to theories of population regulation and helps to identify key factors that have promoted the evolution, maintenance and breakdown of mutualism. I discuss how these ideas might apply to and be tested in ant-plant, fig-wasp and yucca-moth mutualisms.     

Transients: the key to long-term ecological understanding?

Ecological theory has been dominated by a focus on long-term or asymptotic behavior as a way to understand natural systems. Yet experiments are done on much shorter timescales, and the relevant timescales for ecological systems can also be relatively short. Thus, there is a mismatch between the timescales of most experiments and the timescales of many theoretical investigations. However, recent work has emphasized the importance of transient dynamics rather than long-term behavior in ecological systems, enabling the examination of forces that allow coexistence on ecological timescales. Through an examination of what leads to transients in ecological systems, a deeper appreciation of the forces leading to persistence or coexistence in ecological systems emerges, as well as a general understanding of how population levels can change through time.     

生态位分化与森林群落物种多样性维持研究展望

一直以来,生态学家和进化生物学家对森林群落物种多样格局及其形成机制持有不同的观点。虽然Robert Ricklefs将进化和生态过程整合的观点已经被群落生态学家广泛接受,但是区域物种进化历史以及局域群落微进化过程是否能够影响群落生态学过程以及这些过程如何影响群落结构和动态还有待商榷。经典的生态位理论同时强调了种间和种内生态位分化对群落多样性维持的影响。但是生态学家普遍认为种间差异足以代表群落内个体间的相互作用关系,并且由于进化过程导致的种内分化往往涉及较长的时间尺度,因此,虽然种内差异是自然选择的重要材料,物种对环境的适应性进化过程所导致的种内分化对群落构建的影响往往被生态学家所忽视。为此,通过回顾种间和个体生态位分化的研究历史,对两类研究分别进行简要阐述,强调在今后的群落生态学研究中需要考虑个体分化对局域群落构建的影响。     

Not enough niches: non‐equilibrial processes promoting species coexistence in diverse ant communities

Abstract Explanations for species coexistence in ant communities have traditionally focused on niche partitioning, particularly relating to differences in diet, foraging times and nesting requirements. Although niche separation is undoubtedly important, it seems insufficient to account for the high levels of local species richness that are commonly observed. This paper explores alternative explanations for ant species coexistence, focusing on factors that prevent competitive exclusion in diverse ant communities experiencing high levels of behavioural dominance, such as characteristically occurs in Australia. Very high species densities require two conditions to be met: first, a large number of species must successfully establish; and second, there must be a high rate of species persistence once established. In this context I advance five propositions based around three sets of arguments. First, ant sociality and modularity confers a high level of persistence once colonies are established, so that species coexistence is determined to a significant extent by processes operating at the establishment phase, rather than just by interactions between established colonies. Second, competitive outcomes are highly conditioned by environmental variation, which severely limits competitive exclusion. Finally, dominant species are highly patchy in space, and cannot comprehensively monopolize resources, such that there will usually be room for low densities of subordinate species. These propositions have relevance to neutral theories of community ecology, and to understanding intercontinental differences in local species richness.     

The role of variability and risk on the persistence of shared-enemy,predator-prey assemblages

The role of indirect effects such as apparent competition in structuring predator-prey assemblages has recently received empirical attention. That one prey species can be excluded by the impact of a shared-enemy contrasts with the known diversity of multispecies predator-prey interactions. Here, the role of predator foraging among patches of two different prey species is examined as a mechanism that can mediate coexistence in multispecies prey-predator assemblages. Specifically, models of host-parasitoid interactions are constructed to analyse how different types of aggregative behaviour (generated by host-dependent and host-independent responses) affect persistence of the assemblage. How the distribution of hosts and the response of the parasitoid to these distributions can influence coexistence is shown. A generic explanation for coexistence suggests that it is the variability rather than the precise functional relationship that is critical for coexistence under shared-enemy interactions.     

Starve a competitor: evolution of luxury consumption as a competitive strategy

Organisms are often observed to acquire an excess of non-limiting resources, a process known as luxury consumption. Luxury consumption has been largely treated as a bet hedging strategy for temporal variation in resource supply, but may also function as a competitive strategy. We incorporate luxury resource consumption into a derivation of the classic resource ratio model for competition between terrestrial plant, and explore its consequences for population dynamics and competition. We show that luxury consumption reduces the potential for coexistence between two species competing for two resources. Furthermore, we demonstrate that luxury consumption can be selected for because of the competitive advantage that luxury consumers gain. Luxury consumption evolves when competition for resources is local rather than global, there is potential for coexistence between the two species and the competitive environment remains stable over a sufficient period of time to allow selection to act. The evolutionary outcome can be either extinction of one of the competing species or coexistence of the two species with maximum luxury consumption. The potential for selection to favor luxury consumption is well predicted by the competitive outcome between individuals of the two species with and without luxury consumption.     

Spatial Complementarity and the Coexistence of Species

Coexistence of apparently similar species remains an enduring paradox in ecology. Spatial structure has been predicted to enable coexistence even when population-level models predict competitive exclusion if it causes each species to limit its own population more than that of its competitor. Nevertheless, existing hypotheses conflict with regard to whether clustering favours or precludes coexistence. The spatial segregation hypothesis predicts that in clustered populations the frequency of intra-specific interactions will be increased, causing each species to be self-limiting. Alternatively, individuals of the same species might compete over greater distances, known as heteromyopia, breaking down clusters and opening space for a second species to invade. In this study we create an individual-based model in homogeneous two-dimensional space for two putative sessile species differing only in their demographic rates and the range and strength of their competitive interactions. We fully characterise the parameter space within which coexistence occurs beyond population-level predictions, thereby revealing a region of coexistence generated by a previously-unrecognised process which we term the triadic mechanism. Here coexistence occurs due to the ability of a second generation of offspring of the rarer species to escape competition from their ancestors. We diagnose the conditions under which each of three spatial coexistence mechanisms operates and their characteristic spatial signatures. Deriving insights from a novel metric — ecological pressure — we demonstrate that coexistence is not solely determined by features of the numerically-dominant species. This results in a common framework for predicting, given any pair of species and knowledge of the relevant parameters, whether they will coexist, the mechanism by which they will do so, and the resultant spatial pattern of the community. Spatial coexistence arises from complementary combinations of traits in each species rather than solely through self-limitation.     

Trait plasticity alters the range of possible coexistence conditions in a competition–colonisation trade‐off

Most of the classical theory on species coexistence has been based on species‐level competitive trade‐offs. However, it is becoming apparent that plant species display high levels of trait plasticity. The implications of this plasticity are almost completely unknown for most coexistence theory. Here, we model a competition–colonisation trade‐off and incorporate trait plasticity to evaluate its effects on coexistence. Our simulations show that the classic competition–colonisation trade‐off is highly sensitive to environmental circumstances, and coexistence only occurs in narrow ranges of conditions. The inclusion of plasticity, which allows shifts in competitive hierarchies across the landscape, leads to coexistence across a much broader range of competitive and environmental conditions including disturbance levels, the magnitude of competitive differences between species, and landscape spatial patterning. Plasticity also increases the number of species that persist in simulations of multispecies assemblages. Plasticity may generally increase the robustness of coexistence mechanisms and be an important component of scaling coexistence theory to higher diversity communities.     

Allelopathic adaptation can cause competitive coexistence

The maintenance of plant diversity is often explained by the ecological and evolutionary consequences of resource competition. Recently, the importance of allelopathy for competitive interactions has been recognized. In spite of such interest in allelopathy, we have few theories for understanding how the allelopathy influences the ecological and evolutionary dynamics of competing species. Here, I study the coevolutionary dynamics of two competing species with allelopathy in an interspecific competition system, and show that adaptive trait dynamics can cause cyclic coexistence. In addition, very fast adaptation such as phenotypic plasticity is likely to stabilize the population cycles. The results suggest that adaptive changes in allelopathy can lead to cyclic coexistence of plant species even when their ecological characters are very similar and interspecific competition is stronger than intraspecific competition, which should destroy competitive coexistence in the absence of adaptation.     

How competitive intransitivity and niche overlap affect spatial coexistence

Competitive intransitivity is mostly considered outside the main body of coexistence theories that rely primarily on the role of niche overlap and differentiation. How the interplay of competitive intransitivity and niche overlap jointly affects species coexistence has received little attention. Here, we consider a rock–paper–scissors competition system where interactions between species can represent the full spectra of transitive–intransitive continuum and niche overlap/differentiation under different levels of competition asymmetry. By comparing results from pair approximation that only considers interference competition between neighbouring cells in spatial lattices, with those under the mean-field assumption, we show that 1) species coexistence under transitive competition is only possible at high niche differentiation; 2) in communities with partial or pure intransitive interactions, high levels of niche overlap are not necessary to beget species extinction; and 3) strong spatial clustering can widen the condition for intransitive loops to facilitate species coexistence. The two mechanisms, competitive intransitivity and niche differentiation, can support species persistence and coexistence, either separately or in combination. Finally, the contribution of intransitive loops to species coexistence can be enhanced by strong local spatial correlations, modulated and maximised by moderate competition asymmetry. Our study, therefore, provides a bridge to link intransitive competition to other generic ecological theories of species coexistence.     

Feeding strategies,resource utilisation and potential mechanisms for competitive coexistence of Atlantic salmon and alpine bullhead in a sub-Arctic river

We scrutinised the seasonal food-niche utilisation of river dwelling Atlantic salmon parr and alpine bullhead in order to examine potential mechanisms that may facilitate coexistence of species with similar niches. Fish were sampled monthly during the ice-free season, and diet composition and feeding strategy of the two species were compared by analyses of stomach contents. The dietary niches and feeding strategy of salmon parr and bullheads were highly similar both at the individual and population levels, with high within-phenotype contributions to niche width and pronounced generalisation observed during time periods with severe resource limitations. Our findings suggest that competitive coexistence with similar niches may be facilitated by a generalisation of niche width as predicted by optimal foraging theory, rather than the specialised niche width predicted by classic niche theory as a response to interspecific competition. Competitive coexistence may be particularly widespread in spatially and temporally dynamic habitats such as northern lotic systems, which thus may select for generalisation and convergence of ecological niches.     

Trait-based plant ecology: moving towards a unifying species coexistence theory

Functional traits are the center of recent attempts to unify key ecological theories on species coexistence and assembling in populations and communities. While the plethora of studies on the role of functional traits to explain patterns and dynamics of communities has rendered a complex picture due to the idiosyncrasies of each study system and approach, there is increasing evidence on their actual relevance when aspects such as different spatial scales, intraspecific variability and demography are considered.     

Neutral theory and community ecology

I review the mathematical and biological aspects of Hubbell's (2001) neutral theory of species abundance for ecological communities, and clarify its historical connections with closely related approaches in population genetics. A selective overview of the empirical evidence for and against this theory is provided, with a special emphasis on tropical plant communities. The neutral theory predicts many of the basic patterns of biodiversity, confirming its heuristic power. The strict assumption of equivalence that defines neutrality, equivalence among individuals, finds little empirical support in general. However, a weaker assumption holds for stable communities, the equivalence of average fitness among species. One reason for the surprising success of the neutral theory is that all the theories of species coexistence satisfying the fitness equivalence assumption, including many theories of niche differentiation, generate exactly the same patterns as the neutral theory. Hubbell's neutral theory represents an important synthesis and a much needed demonstration of the pivotal role of intraspecific variability in ecology. Further improvements should lead to an explicit linking to niche‐based processes. This research programme will depend crucially on forthcoming theoretical and empirical achievements.     

Elevated carbon dioxide is predicted to promote coexistence among competing species in a trait‐based model

Differential species responses to atmospheric CO₂ concentration (C_a) could lead to quantitative changes in competition among species and community composition, with flow‐on effects for ecosystem function. However, there has been little theoretical analysis of how elevated C_a (eC_a) will affect plant competition, or how composition of plant communities might change. Such theoretical analysis is needed for developing testable hypotheses to frame experimental research. Here, we investigated theoretically how plant competition might change under eC_a by implementing two alternative competition theories, resource use theory and resource capture theory, in a plant carbon and nitrogen cycling model. The model makes several novel predictions for the impact of eC_a on plant community composition. Using resource use theory, the model predicts that eC_a is unlikely to change species dominance in competition, but is likely to increase coexistence among species. Using resource capture theory, the model predicts that eC_a may increase community evenness. Collectively, both theories suggest that eC_a will favor coexistence and hence that species diversity should increase with eC_a. Our theoretical analysis leads to a novel hypothesis for the impact of eC_a on plant community composition. This hypothesis has potential to help guide the design and interpretation of eC_a experiments.     

The present status of the competitive exclusion principle

Since Darwin accepted the Malthusian population theory to solve the demographic problems he thought to be logically connected with the universal operation of natural selection, the numerical processes in both populations and communities were generally supposed to be governed by competition. For interspecific relations this found expression in the 'competitive exclusion principle'. After it was shown that coexistence rather than exclusion of closely related species is the rule, this principle gradually changed into the 'competitive niche shift principle'. Recently the universality of competition has been increasingly questioned, so that other interspecific relationships (especially predation) are revaluated as possibly governing many natural population and inter-population processes.