Coexistence of fish species with strongly similar life histories - population dynamical feedback forces species to pick sides

Fish generally grow several orders of magnitude between the larval and adult stage. Many ecological properties of organisms are related to body size, and hence small fish often have very different ecological roles than large conspecifics. This also implies that omnivory, the feeding on more than one trophic level by individuals of the same species, is a common phenomenon in fish. Intraguild predation is omnivory in its simplest form, where two species compete for the same resource, but one of the species can also eat its competitor. In models, persistence of both species in such a configuration is difficult to obtain. In marine fish communities however, it is observed routinely. One way in which persistence of omnivorous species can be established is by incorporating it as an ontogenetic diet shift, where small individuals of both species compete, and large individuals of one can feed on the small individuals of the others species. We show in this study that this mechanism can not only lead to persistence of a single omnivorous species, but also to persistence of multiple omnivorous species. This is possible given that the adults have sufficiently different diets. It is shown that, while adults of both species can feed both on small competitors or on the basic resource, due to the population dynamical feedback, one species will in practice act as a predator, while the other acts as prey. This way, a system with two omnivores in practice persists as a tritrophic system. Which of the species assumes which role depends on the specific community characteristics. We show here that by incorporating complex size-dependent feeding relationships in food webs, many more species may be able to coexist than is possible based on either species-level considerations or size spectrum models which do incorporate within-population size differences, but relate diet only to individual body size irrespective of species identity. The mechanism underlying our result may be part of the explanation why fish species with seemingly similar life histories do coexist in marine ecosystems

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

Investigations into stability in the fig/ fig-wasp mutualism

Fig trees (Ficus, Moraceae) and their pollinating wasps (Chalcidoidea, Agaonidae) are involved in an obligate mutualism where each partner relies on the other in order to reproduce: the pollinating fig wasps are a fig tree’s only pollen disperser whilst the fig trees provide the wasps with places in which to lay their eggs. Mutualistic interactions are, however, ultimately genetically selfish and as such, are often rife with conflict. Fig trees are either monoecious, where wasps and seeds develop together within fig fruit (syconia), or dioecious, where wasps and seeds develop separately. In interactions between monoecious fig trees and their pollinating wasps, there are conflicts of interest over the relative allocation of fig flowers to wasp and seed development. Although fig trees reap the rewards associated with wasp and seed production (through pollen and seed dispersal respectively), pollinators only benefit directly from flowers that nurture the development of wasp larvae, and increase their fitness by attempting to oviposit in as many ovules as possible. If successful, this oviposition strategy would eventually destroy the mutualism; however, the interaction has lasted for over 60 million years suggesting that mechanisms must be in place to limit wasp oviposition. This thesis addresses a number of factors to elucidate how stability may be achieved in monoecious fig systems. Possible mechanisms include: 1) a parasitoidcentred short ovipositor hypothesis in Ficus rubiginosa, which suggests that a subset of flowers are out of reach to parasitoid ovipositors making these ovules the preferred choice for ovipositing pollinators and allowing seeds to develop in less preferred ovules; 2) the presence of third-party mutualists such as non-pollinating fig wasps (F. burkei) and patrolling green tree ants on the fig surface (F. racemosa) that limit pollinator and parasitoid oviposition respectively; and 3) selection on fig morphology which constrains the size (and therefore fecundity) of the associated pollinators. I discuss the lack of evidence for a single unifying theory for mutualism stability and suggest that a more likely scenario is the presence of separate, and perhaps multiple, stabilising strategies in different fig/ fig-wasp partnerships

The effect of canopy position on growth and mortality in mixed sapling communities during self-thinning

This research investigates how species in the sapling phase differ in growth and survival depending on light availability (as estimated by canopy position) by means of tree-ring analysis and modelling mortality. We harvested 120 live and 158 dead saplings in self-thinning communities consisting of Silver birch (Betula pendula Roth.), Scots pine (Pinus sylvestris L.), Japanese larch (Larix kaempferi Carr.) and Douglas fir (Pseudotsuga menziesii Mirb. Franco) in the Netherlands. Results are evaluated within the framework of a trade-off between high-light growth and low-growth survival. Radial growth, measured at ground level, generally declined over time. In addition, a decreasing light availability further reduced growth in all species except Douglas fir. Trees died when radial growth was reduced to about 0.5 mm year¿1. Mortality in all species except Scots pine was significantly related to recent growth, but mortality curves were not different. The light-demanding Silver birch and Japanese larch differed from the shade-tolerant Douglas fir in both high-light growth and low-growth mortality, in line with a growth-survival trade-off. The light-demanding Scots pine did not fit this pattern as it was unable to transfer high radial growth into height gain, leaving it in suppressed canopy positions. This indicates the importance of height growth in the growth-survival trade-off. Differences in mortality probabilities affect the potential for coexistence, however, in all species also fast-growing individuals died suggesting additional factors causing mortality during self-thinning, other than direct competition for ligh

Robust permanence for ecological equations with internal and external feedbacks.

Species experience both internal feedbacks with endogenous factors such as trait evolution and external feedbacks with exogenous factors such as weather. These feedbacks can play an important role in determining whether populations persist or communities of species coexist. To provide a general mathematical framework for studying these effects, we develop a theorem for coexistence for ecological models accounting for internal and external feedbacks. Specifically, we use average Lyapunov functions and Morse decompositions to develop sufficient and necessary conditions for robust permanence, a form of coexistence robust to large perturbations of the population densities and small structural perturbations of the models. We illustrate how our results can be applied to verify permanence in non-autonomous models, structured population models, including those with frequency-dependent feedbacks, and models of eco-evolutionary dynamics. In these applications, we discuss how our results relate to previous results for models with particular types of feedbacks

Daisyworld: a review

Daisyworld is a simple planetary model designed to show the long-term effects of coupling between life and its environment. Its original form was introduced by James Lovelock as a defense against criticism that his Gaia theory of the Earth as a self-regulating homeostatic system requires teleological control rather than being an emergent property. The central premise, that living organisms can have major effects on the climate system, is no longer controversial. The Daisyworld model has attracted considerable interest from the scientific community and has now established itself as a model independent of, but still related to, the Gaia theory. Used widely as both a teaching tool and as a basis for more complex studies of feedback systems, it has also become an important paradigm for the understanding of the role of biotic components when modeling the Earth system. This paper collects the accumulated knowledge from the study of Daisyworld and provides the reader with a concise account of its important properties. We emphasize the increasing amount of exact analytic work on Daisyworld and are able to bring together and summarize these results from different systems for the first time. We conclude by suggesting what a more general model of life-environment interaction should be based on

The complex net effect of reciprocal interactions and recruitment facilitation maintains an intertidal kelp community

1. Theoretical and empirical ecology has transitioned from a focus on the role of negative interactions in species coexistence to a more pluralistic view that acknowledges that coexistence in natural communities is more complex, and depends on species interactions that vary in strength, sign, and reciprocity, and such contexts as the environment and life-history stage. 2. We used a whole-community approach to examine how species interactions contribute to the maintenance of a rocky intertidal macroalgal canopy–understorey assemblage. We determined both the types of interactions in this network, and whether interactions were sensitive to environmental gradients. 3. Focusing on a structurally dominant canopy kelp Saccharina sessilis, and its diverse co-occurring understorey assemblage, we evaluated the role of the understorey in controlling S. sessilis recruitment and quantified the reciprocal effect of the S. sessilis canopy and understorey on one another using a removal experiment replicated across 600 km of coastline. We determined the sensitivity of interactions to natural variation in light and nutrient availability (replicated among four regions on the N.E. Pacific coast), and under different wave conditions (three wave regimes). 4. Surprisingly, species interactions were similar across sites and thus not context-dependent. Unexpectedly, the understorey community had a strong positive effect on the S. sessilis canopy, whereby the adult canopy decreased dramatically following understorey removal. Additionally, S. sessilis recruitment depended on the presence of understorey coralline algal turf. In turn, the canopy had a neutral effect on the coralline understorey, but a negative effect on non-calcifying algal turfs, likely eventually generating positive indirect canopy effects on the coralline understorey. Density-dependent intraspecific competition between S. sessilis adults and recruits may moderate this positive feedback between the S. sessilis canopy and coralline understorey. 5. Synthesis. Our research highlights the importance of positive interactions for coexistence in natural communities, and the necessity of studying multiple life-history stages and reciprocal species interactions in order to elucidate the mechanisms that maintain diversity.Keywords: marine, plant-plant interactions, aquatic plant ecology, positive interactions, environmental gradients, benthic, macroalga