Macroevolutionary patterns in the origin of mutualisms involving ants

Ants are a diverse and abundant insect group that form mutualistic associations with a number of different organisms from fungi to insects and plants. Here, we use a phylogenetic approach to identify ecological factors that explain macroevolutionary trends in the mutualism between ants and honeydew-producing Homoptera. We also consider association between ant-Homoptera, ant-fungi and ant-plant mutualisms. Homoptera-tending ants are more likely to be forest dwelling, polygynous, ecologically dominant and arboreal nesting with large colonies of 10(4)-10(5) individuals. Mutualistic ants (including those that garden fungi and inhabit ant-plants) are found in under half of the formicid subfamilies. At the genus level, however, we find a negative association between ant-Homoptera and ant-fungi mutualisms, whereas there is a positive association between ant-Homoptera and ant-plant mutualisms. We suggest that species can only specialize in multiple mutualisms simultaneously when there is no trade-off in requirements from the different partners and no redundancy of rewards.     

Asymmetric or diffusive co-evolution generates meta-populations in fig-fig wasp mutualisms

Co-evolutionary theory assumes co-adapted characteristics are a positive response to counter those of another species,whereby co-evolved species reach an evolutionarily stable interaction through bilateral adaptation.However,evidence from the fig-fig wasp mutualistic system implies very different co-evolutionary selection mechanisms,due to the inherent conflict among interacted partners.Fig plants appear to have discriminatively enforced fig wasps to evolve"adaptation characteristics"that provide greater benefit to the fig,and fig wasps appear to have diversified their evolutionary strategies in response to discriminative enforcement by figs and competition among different fig wasp species.In what appears to be an asymmetric interaction,the prosperity of cooperative pollinating wasps should inevitably lead to population increases of parasitic individuals,thus resulting in localized extinctions of pollinating wasps.In response,the sanctioning of parasitic wasps by the fig should lead to a reduction in the parasitic wasp population.The meta-populations created by such asymmetric interactions may result in each population of coevolved species chaotically oscillated,temporally or evolutionarily.     

A general model for the evolution of mutualisms

The evolution of mutualisms presents a puzzle. Why does selection favour cooperation among species rather than cheaters that accept benefits but provide nothing in return? Here we present a general model that predicts three key factors will be important in mutualism evolution: (i) high benefit to cost ratio, (ii) high within‐species relatedness and (iii) high between‐species fidelity. These factors operate by moderating three types of feedback benefit from mutualism: cooperator association, partner‐fidelity feedback and partner choice. In defining the relationship between these processes, our model also allows an assessment of their relative importance. Importantly, the model suggests that phenotypic feedbacks (partner‐fidelity feedback, partner choice) are a more important explanation for between‐species cooperation than the development of genetic correlations among species (cooperator association). We explain the relationship of our model to existing theories and discuss the empirical evidence for our predictions.     

Foundress re-emergence and fig permeability in fig tree-wasp mutualisms

Some female pollinating fig wasps (foundresses) re-emerge from figs after oviposition/pollination. We investigated why this occurs in the mutualism between the gynodioecious Ficus montana and Liporrhopalum tentacularis. Re-emergence increased with foundress density in figs and some foundresses oviposited in two male figs, indicating that they re-emerge because of oviposition site limitation. Re-emergence was independent of fig diameter, indicating that permeability is not because of fig age at entry. Rather, as some foundresses also pollinate two female figs we suggest permeability is selected for because it increases pollinator production and/or efficiency (although, potentially opposing these hypotheses, we also found between-tree differences in permeability in male figs). In addition, we show that re-emergence is much more common than previously suspected, and more common from gynodioecious than monoecious fig species. We argue that our findings in F. montana could explain this pattern of incidence.     

The macroecology of marine cleaning mutualisms

1. Marine cleaning mutualisms generally involve small fish or shrimps removing ectoparasites and other material from cooperating 'client' fish. We evaluate the role of fish abundance, body size and behaviour as determinants of interactions with cleaning mutualists. 2. Data come from eight reef locations in Brazil, the Caribbean, the Mediterranean and Australia. 3. We conducted a meta-analysis of client-cleaner interactions involving 11 cleaner and 221 client species. 4. There was a strong, positive effect of client abundance on cleaning frequency, but only a weak, negative effect of client body size. These effects were modulated by client trophic group and social behaviour. 5. This study adds to a growing body of evidence suggesting a central role of species abundance in structuring species interactions.     

Is the cask of facilitation ready for bottling? A symposium on the connectivity and future directions of positive plant interactions

The 2009 British Ecological Society''s Annual Symposium entitled ‘Facilitation in Plant Communities’ was held at the University of Aberdeen, Scotland, from 20 to 22 April 2009. This was the first ever international meeting dedicated to the rapidly expanding field of facilitation. The aim of the symposium was to assess the current ‘state-of-play’ by contrasting findings from different systems and by looking outwards in an attempt to integrate this field with other related fields. It was also aimed at understanding how knowledge of facilitation can help understand community dynamics and be applied to ecosystem restoration. The symposium identified several key areas where future work is likely to be most profitable.     

Allee dynamics generated by protection mutualisms can drive oscillations in trophic cascades

Understanding the relative effect of top predators and primary producers on intermediate trophic levels is a key question in ecology. Most previous work, however, has not considered either realistic nonlinearities in feedback between trophic levels or the effect of mutualists on trophic cascades. Here, we develop a realistic model for a protection mutualism that explicitly includes interactions between a protected herbivore and both its food plant and generalist predators. In the absence of protection, herbivores and plant resources approach a stable equilibrium, provided that predation is not so high as to cause herbivore extinction. In contrast, adding protection by mutualists increases the range of dynamical outcomes to include unstable equilibria, stable and unstable limit cycles, and heteroclinic orbits. By reducing the impact of predators, protection by mutualists can allow herbivores to exert strong negative effects on their host plants, which in turn can lead to repeated cycles of overexploitation and recovery. Our results indicate that it may be essential to consider protection mutualisms to understand the dynamics of trophic cascades. Conversely, it may be essential to explicitly include dynamical feedback between plants and herbivores to fully understand the population and community dynamical consequences of protection mutualism.     

Ant specificity and behaviour in mutualisms with epiphytes: the case of Lecanopteris (Polypodiaceae)

Seven species of the fern Lecanopteris: L. sinuosa, L. sarcopus, L. mirabilis, L. curtisii, L. pumila, L. celebica and L. damaedii are regularly inhabited by ants of five species: Iridomyrmex cordatus, I. murinus, Crematogaster treubi, C. difformis and Camponotus pallidas. Inhabitation is not obligate; either party can survive without the other, but ferns without ants are rare in their natural habitat. The total recorded diversity of ants in Lecanopteris is 31 species, 20 of which were only recorded once.
All five regularly inhabiting ant species kept larvae in the domatium (82% of Lecanopteris specimens examined). The major ant species inhabiting Lecanopteris deposited debris in the domatium, segregated from their brood (90% of Lecanopteris specimens). Iridomyrmex murinus did not build carton around the rhizomes, but the other major inhabitants constructed runways in 79% of Lecanopteris specimens. Goccid cultivation was variable within ant and plant species: I murinus tended them in 50% of specimens, and homopterans were also recorded with Crematogaster treubi (39%), C. bomeensis (27%) and I cordatus (16%). A high frequency of ant colonies inhabiting Lecanopteris , keeping larvae, depositing debris, building carton runways, and occasionally keeping coccids has been established. Specificity of ant species is high within a population of Lecanopteris , or a given habitat or geographical area, but outside the ranges of regular inhabitants other ant occupants are found. Comparable data from other genera of ant-epiphytes with domatia show similar diversity of inhabitance over geographical areas; no data are available within single plant populations.