Colony structure in a plant-ant: behavioural,chemical and genetic study of polydomy in<Emphasis Type="Italic"> Cataulacus mckeyi</Emphasis> (Myrmicinae)

Social organisation of colonies of obligate plant-ants can affect their interaction with myrmecophyte hosts and with other ants competing for the resources they offer. An important parameter of social organisation is whether nest sites of a colony include one or several host individuals. We determined colony boundaries in a plant-ant associated with the rainforest understorey tree Leonardoxa africana subsp. africana, found in coastal forests of Cameroon (Central Africa). This myrmecophyte is strictly associated with two ants, Petalomyrmex phylax and Cataulacus mckeyi. Plants provide food and nesting sites for P. phylax, which protects young leaves against insect herbivores. This mutualism is often parasitised by C. mckeyi, which uses but does not protect the host. The presence of C. mckeyi on a tree excludes the mutualistic ant. Because Petalomyrmex -occupied trees are better protected, their growth and survival are superior to those of Cataulacus -occupied trees, giving P. phylax an advantage in occupation of nest sites. C. mckeyi often colonises trees that have lost their initial associate P. phylax, as a result of injury to the tree caused by disturbance. Polydomy may allow C. mckeyi to occupy small clumps of trees, without the necessity of claustral colony foundation in each tree. Investigating both the proximate (behavioural repertoire, colony odour) and the ultimate factors (genetic structure) that may influence colony closure, we precisely defined colony boundaries. We show that colonies of C. mckeyi are monogynous and facultatively polydomous, i.e. a colony occupies one to several Leonardoxa trees. Workers do not produce males. Thus, the hypothesis that polydomy allows workers in queenless nests to evade queen control for their reproduction is not supported in this instance. This particular colony structure may confer on C. mckeyi an advantage in short-distance dispersal, and this could help explain its persistence within the dynamic Leonardoxa system.     

Perspective: sexual conflict and sexual selection: chasing away paradigm shifts

Abstract.— Traditional models of sexual selection propose that partner choice increases both average male and average female fitness in a population. Recent theoretical and empirical work, however, has stressed that sexual conflict may be a potent broker of sexual selection. When the fitness interests of males and females diverge, a reproductive strategy that increases the fitness of one sex may decrease the fitness of the other sex. The chase-away hypothesis proposes that sexual conflict promotes sexually antagonistic, rather than mutualistic, coevolution, whereby manipulative reproductive strategies in one sex are counteracted by the evolution of resistance to such strategies in the other sex. In this paper, we consider the criteria necessary to demonstrate the chase-away hypothesis. Specifically, we review sexual conflict with particular emphasis on the chase-away hypothesis; discuss the problems associated with testing the predictions of the chase-away hypothesis and the extent to which these predictions and the predictions of traditional models of sexual selection are mutually exclusive; discuss misconceptions and mismeasures of sexual conflict; and suggest an alternative approach to demonstrate sexual conflict, measure the intensity of sexually antagonistic selection in a population, and elucidate the coevolutionary trajectories of the sexes.     

Mutualistic relationships between phytoplankton and bacteria caused by carbon excretion from phytoplankton

For the competition system of phytoplankton and bacteria through inorganic phosphorus, our mathematical model showed that mutualistic relationships between them could occur due to production and consumption of extracellular organic carbon by phytoplankton and bacteria. In our model, phytoplankton are limited in their growth by light and phosphorus, and bacteria are limited in their growth by phosphorus and carbon released from phytoplankton. We adopted permanence as a criterion of the coexistence in mathematical analysis, and led necessary conditions of permanence in the model. Under these coexistence conditions, we estimated the strength of total effects of interactions between phytoplankton and bacteria at the steady state by press perturbation method. The results of this estimation indicated the mutualistic interactions between phytoplankton and bacteria. This suggests that mutualistic situation could occur due to the introduction of carbon flow from phytoplankton to bacteria, even if phytoplankton and bacteria compete with each other through common resource, inorganic phosphorus.     

Speciation in fig pollinators and parasites

Here we draw on phylogenies of figs and fig wasps to suggest how modes of speciation may be affected by interspecific interactions. Mutualists appear to have cospeciated with their hosts to a greater extent than parasites, which showed evidence of host shifting. However, we also repeatedly encountered a pattern not explained by either cospeciation or host switching. Sister species of fig parasites often attack the same host in sympatry, and differences in ovipositor length suggest that parasite speciation could result from divergence in the timing of oviposition with respect to fig development. These observations on fig parasites are consistent with a neglected model of sympatric speciation.     

Age-specific patterns in honeydew production and honeydew composition in the aphid Metopeurum fuscoviride: implications for ant-attendance

The intensity of the mutualistic relationship between aphids and ants depends mainly on the composition and amount of honeydew. We used the model system Tanacetum vulgare-Metopeurum fuscoviride to study age-related differences in honeydew production and composition and its effect on the mutualism between M. fuscoviride and the ant Lasius niger. First and second instar larvae of M. fuscoviride produced only half of the amount of honeydew as older larvae or adults. There were, however, no differences between age classes in the total honeydew sugar concentration, which averaged approx. 80 μg sugar/μl honeydew. Honeydew sugar composition also did not differ between age classes, and melezitose was the dominant sugar (59% in all classes). The amino acid concentration, by contrast, increased significantly with aphid age, reaching 22.6 nmol per μl honeydew in adult M. fuscoviride. This increase was mainly caused by asparagine and glutamine, while there were no differences in the concentrations of the five other regularly detected amino acids and cystine, respectively. The intensity of ant-attendance was significantly lower in colonies of first and second instar larvae than in colonies of older age classes. Ant-attendance correlated with the amount of honeydew produced, and not with the total amino acid concentration.     

GRAZING OPTIMIZATION,NUTRIENT CYCLING,AND SPATIAL HETEROGENEITY OF PLANT‐HERBIVORE INTERACTIONS: SHOULD A PALATABLE PLANT EVOLVE?

Abstract.— Can the evolution of plant defense lead to an optimal primary production? In a general theoretical model, Loreau (1995) and de Mazancourt et al. (1998, 1999) have shown that herbivory could increase primary production up to a moderate rate of grazing intensity through recycling of a limiting nutrient, provided several conditions are fulfilled. In the present paper, we assume: (1) grazing intensity is controlled by plants through their level of palatability; and (2) plant fitness is determined by its productivity. We explore the conditions under which such an optimal production may be reached through natural selection. We model two competing plant types that differ only in palatability and are distributed in a patchy landscape determined by the plant‐herbivore interaction. Patch size is determined by herbivore behavior: herbivores recycle nutrient homogeneously within patches, but recycle nutrient proportionally to consumption between patches. The model shows that a strategy of intermediate palatability can be adaptive in response to a small herbivore that lives on and recycles nutrient around one or a few individual plants. For moderately small herbivores, plant palatability may evolve towards one of two local convergent strategies, depending on the initial conditions. For medium‐ to large‐sized herbivores, the nonpalatable strategy is always selected. We discuss the functional and evolutionary implications of these results, and suggest that the traditional dichotomy describing antagonistic and mutualistic interactions may be misleading.     

Do submerged aquatic plants influence periphyton community composition for the benefit of invertebrate mutualists?

• 1 It has been suggested that submerged aquatic plants can influence the periphyton which grows on their surfaces, making it nutritionally beneficial to snails. In return, preferential feeding by snails clears the plants from a potential competitor, with both plants and grazers gaining from this mutualistic relationship.
• 2 A highly replicated experiment was conducted, in which the nature of the plant (isoetid and elodeid types compared with similar shaped inert substrata), the nutrient availability (10–200 µg L^‐1 P, 0.2–4 mg L^‐1 N) and the influence of periphyton grazers, Physa fontinalis, were controlled. The plants were cleaned of periphyton before use and an algal inoculum added to all treatments. At the end of the growth period, quantitative measures of the periphyton community composition were made and related to the treatments using both ordination and analysis of variance.
• 3 Grazing had the largest influence on community composition and algal numbers. A community of unicellular and adpressed filamentous forms developed in the presence of snails, and of erect filamentous forms in their absence. Three algal species, Cocconeis placentula, Chamaesiphon incrustans and Aphanochaete repens, increased in real numbers in the presence of snails, probably as a result of reduced competition whilst being able to withstand grazing.
• 4 The second largest effect was the influence of host plant. However, differences between the two artificial plants were as great as between the real plants and their artificial counterparts, indicating that physical structure was as important as any active contribution by the plants. Nutrients had a small but significant effect on community composition, but not all species responded in the same way to nutrient enrichment.
• 5 Although submerged aquatic plants exert an influence over the community composition of the periphyton which develops on their surfaces, it is unlikely that they manipulate it to make it more attractive to grazers such as snails.

     

Leaf domatia and foliar mite abundance in broadleaf deciduous forest of north Asia

Plant morphology may be shaped, in part, by the third trophic level. Leaf domatia, minute enclosures usually in vein axils on the leaf underside, may provide the basis for protective mutualism between plants and mites. Domatia are particularly frequent among species of trees, shrubs, and vines in the temperate broadleaf deciduous forests in north Asia where they may be important in determining the distribution and abundance of mites in the forest canopy. In lowland and montane broadleaf deciduous forests at Kwangn;akung and Chumbongsan in Korea, we found that approximately half of all woody species in all forest strata, including many dominant trees, have leaf domatia. Pooling across 24 plant species at the two sites, mites occupied a mode of 60% (range 20-100%) of domatia and used them for shelter, egg-laying, and development. On average, 70% of all active mites and 85% of mite eggs on leaves were found in domatia; over three-quarters of these were potentially beneficial to their hosts. Further, mite abundance and reproduction (expressed as the proportion of mites at the egg stage) were significantly greater on leaves of species with domatia than those without domatia in both forests. Effects of domatia on mite abundance were significant only for predaceous and fungivorous mite taxa; herbivore numbers did not differ significantly between leaves of species with and without domatia. Comparable patterns in broadleaf deciduous forest in North America and other biogeographic regions suggest that the effect of leaf domatia on foliar mite abundance is general. These results are consistent with several predictions of mutualism between plants and mites, and indicate that protective mutualisms may be frequent in the temperate zone.     

THE EVOLUTION OF HOST-PLANT USE AND SEQUESTRATION IN THE LEAF BEETLE GENUS PHRATORA (COLEOPTERA: CHRYSOMELIDAE)

Leaf beetles in the genus Phratora differ in host plant use and in the chemical composition of their larval defensive secretion. Most species specialize on either poplars or willows (family Salicaceae), but two species feed on birch (family Betulaceae). Phratora vitellinae utilizes salicylates from the host plant to produce its larval secretion, which contains salicylaldehyde, while other Phratora species produce an autogenous secretion. To reconstruct the evolutionary history of host plant use and the larval secretion chemistry in this genus, we sequenced 1383 base pairs of the mt cytochrome oxidase I gene for six European and one North American Phratora species and three outgroup taxa. Bootstrap values of the complete nucleotide sequence were 99-100% for six of eight nodes in the maximum parsimony tree. They were 71% and 77% for the two other nodes. The maximum parsimony tree and the maximum likelihood tree based on nucleotide sequence showed the same relationships as a maximum parsimony tree based on the amino acid sequence. Beetle phylogeny overlapped broadly with host plant taxonomy and chemistry, and it revealed historical constraints influencing host plant use. However, there was one host shift from the willow family (Salicaceae) to the birch family (Betulaceae). The use of host plant phenol glycosides for the larval defensive secretion evolved along the lineage that led to P. vitellinae. Phratora vitellinae feeds on the taxonomically widest range of host plants, which are characterized by moderate to high levels of salicylates. The results support the hypothesis that the use of salicylates for the larval secretion evolved twice independently in chrysomeline leaf beetles.