Anti-predator defence mechanisms in sawfly larvae of Arge (Hymenoptera, Argidae)

Larvae of the sawfly Arge (Hymenoptera, Argidae) are exposed to predators such as ants. Their defence mechanisms, which have been almost unstudied, were investigated by behavioural observations coupled to a morphological approach and by testing the bioactivity of several body parts. Arge larvae raised their abdomen when contacted by Myrmica rubra workers. The ants rarely bit a larva and generally retreated immediately, sometimes without contacting it. Most of those few ants that bit a larva then showed an uncoordinated walk. Crude hemolymph from a common species, A. pagana, was a feeding deterrent towards ants. Hemolymph extracts remained active up to a concentration of 0.8 microg DW extract per microlitre solution, and were more active than integument and gut extracts. We also observed ants paralysed by extracts, especially from the gut. It is likely that this toxicity is due to a polypeptide, lophyrotomin, which is known to occur in A. pullata. Six or seven non-eversible ventro-abdominal glands occurred in all species studied (A. fuscipes, A. nigripes, A. ochropus, A. pagana, A. pullata, A. ustulata). These glands contain volatiles. We consider both types of chemicals to be important in defence, and we propose that the paralysing effect is a common feature among Arge species.     

Vision-based ability of an ant-mimicking jumping spider to discriminate between models,conspecific individuals and prey

Myrmarachne assimilis, an ant-like (myrmecomorphic) jumping spider (Araneae, Salticidae) from the Philippines, is a Batesian mimic of Oecophylla smaragdina, the Asian weaver ant. Salticids are well known for their acute eyesight and the elaborate vision-based display behaviour they adopt during encounters with conspecific individuals, but most salticids are not myrmecomorphic. Despite its unusual morphology, M. assimilis adopts display behaviour during intraspecific interactions that is similar to the display behaviour of more typical salticids. The specificity with which M. assimilis deploys display behaviour is investigated and provides insights into this mimic’s ability to differentiate, by sight alone, between models, conspecific individuals and prey. During each standardized test, an adult M. assimilis female was in a large cage along with a small transparent glass vial, a stimulus animal being enclosed in the vial such that potential optical cues, but not potential chemical cues, were available to the tested M. assimilis individual. Depending on the test, the stimulus animal was another adult M. assimilis female, a house fly (prey) or an ant (Camponotus sp. or O. smaragdina). Only the conspecific female consistently elicited display from M. assimilis, implying that M. assimilis is a Batesian mimic that can, when relying on vision alone, discriminate between conspecific individuals, models and prey. Received 12 June 2006; revised 22 September 2006; accepted 26 September 2006.     

Phylogeny and phylogeography of the Mediterranean species of the parasitic ant genus <Emphasis Type="Italic">Chalepoxenus</Emphasis> and its <Emphasis Type="Italic">Temnothorax</Emphasis> hosts

We analysed the phylogenetic and phylogeographic relationships of four Mediterranean species of the rare slave-making ant genus Chalepoxenus and eleven of its about 20 Temnothorax host species by sequencing the mitochondrial Cytochrome Oxidase I and II genes. Neighbour-Joining, Maximum Parsimony and Bayesian analyses based on 1320 bp indicate that the genus Chalepoxenus constitutes a monophylum. In all three analyses, C. kutteri from Southwest Europe and the workerless, “degenerate slavemaker” C. brunneus from North Africa form a monophyletic group. C. muellerianus and C. tauricus, distributed in Southern Europe and Ukraine, respectively, form a monophylum in the Neighbour-Joining and the Maximum Parsimony analysis. In our limited set of only 11 of several hundred Temnothorax species, T. flavicornis forms the sister group of Chalepoxenus. Our study further indicates paraphyly of the genus Temnothorax with respect to Chalepoxenus. Moreover, the results suggest that speciation in this slave-making genus is possibly caused by the formation of host races as different Chalepoxenus species use different hosts, and some samples seem to cluster by host species rather than by geographical distance. Received 5 September 2006; revised 17 March 2007; accepted 23 March 2007.     

Population genetic signatures of diffuse co-evolution between leaf-cutting ants and their cultivar fungi

Switching of symbiotic partners pervades most mutualisms, despite mechanisms that appear to enforce partner fidelity. To investigate the interplay of forces binding and dissolving mutualistic pairings, we investigated partner fidelity at the population level in the attine ant-fungal cultivar mutualism. The ants and their cultivars exhibit both broad-scale co-evolution, as well as cultivar switching, with short-term symbiont fidelity maintained by vertical transmission of maternal garden inoculates via dispersing queens and by the elimination of alien cultivar strains. Using microsatellite markers, we genotyped cultivar fungi associated with five co-occurring Panamanian attine ant species, representing the two most derived genera, leaf-cutters Atta and Acromyrmex. Despite the presence of mechanisms apparently ensuring the cotransmission of symbiont genotypes, different species and genera of ants sometimes shared identical fungus garden genotypes, indicating widespread cultivar exchange. The cultivar population was largely unstructured with respect to host ant species, with only 10% of the structure in genetic variance being attributable to partitioning among ant species and genera. Furthermore, despite significant genetic and ecological dissimilarity between Atta and Acromyrmex, generic difference accounted for little, if any, variance in cultivar population structure, suggesting that cultivar exchange dwarfs selective forces that may act to create co-adaptive ant-cultivar combinations. Thus, binding forces that appear to enforce host fidelity are relatively weak and pairwise associations between cultivar lineages and ant species have little opportunity for evolutionary persistence. This implicates that mechanisms other than partner fidelity feedback play important roles in stabilizing the leafcutter ant-fungus mutualism over evolutionary time.     

Genetic population structure, queen supersedure and social polymorphism in a social Hymenoptera

In social insects, the emergence of multiple queening is linked to changes in a suite of traits such as the reproductive life span of queens, mating patterns and population structure. We investigated queen turnover, colony longevity, spatial distribution patterns and genetic differentiation in a population of the socially polymorphic ant Formica fusca. Genetic differentiation between the social forms was absent, and mating patterns were similar in the two forms. The spatial distribution of single- and multi-queen colonies indicated an absence of colony reproduction by budding in both colony types. However, the rate of queen supersedure was high in multi-queen colonies and absent in single-queen ones. The social structure of colonies remained stable across years, but colony mortality did not differ between the two social forms. These results imply that differences between social types may appear and persist also in sympatry, and that these differences may occur in some traits, but not others, despite the presence of homogenizing gene flow.     

Stable isotopes reveal links between human food inputs and urban ant diets

The amount of energy consumed within an average city block is an order of magnitude higher than that consumed in any other ecosystem over a similar area. This is driven by human food inputs, but the consequence of these resources for urban animal populations is poorly understood. We investigated the role of human foods in ant diets across an urbanization gradient in Manhattan using carbon and nitrogen stable isotopes. We found that some—but not all—ant species living in Manhattan''s most urbanized habitats had δ¹³C signatures associated with processed human foods. In particular, pavement ants (Tetramorium sp. E) had increased levels of δ¹³C similar to δ¹³C levels in human fast foods. The magnitude of this effect was positively correlated with urbanization. By contrast, we detected no differences in δ¹⁵N, suggesting Tetramorium feeds at the same trophic level despite shifting to human foods. This pattern persisted across the broader ant community; species in traffic islands used human resources more than park species. Our results demonstrate that the degree urban ants exploit human resources changes across the city and among species, and this variation could play a key role in community structure and ecosystem processes where human and animal food webs intersect.     

Evolutionarily advanced ant farmers rear polyploid fungal crops

Innovative evolutionary developments are often related to gene or genome duplications. The crop fungi of attine fungus‐growing ants are suspected to have enhanced genetic variation reminiscent of polyploidy, but this has never been quantified with cytological data and genetic markers. We estimated the number of nuclei per fungal cell for 42 symbionts reared by 14 species of Panamanian fungus‐growing ants. This showed that domesticated symbionts of higher attine ants are polykaryotic with 7–17 nuclei per cell, whereas nonspecialized crops of lower attines are dikaryotic similar to most free‐living basidiomycete fungi. We then investigated how putative higher genetic diversity is distributed across polykaryotic mycelia, using microsatellite loci and evaluating models assuming that all nuclei are either heterogeneously haploid or homogeneously polyploid. Genetic variation in the polykaryotic symbionts of the basal higher attine genera Trachymyrmex and Sericomyrmex was only slightly enhanced, but the evolutionarily derived crop fungi of Atta and Acromyrmex leaf‐cutting ants had much higher genetic variation. Our opposite ploidy models indicated that the symbionts of Trachymyrmex and Sericomyrmex are likely to be lowly and facultatively polyploid (just over two haplotypes on average), whereas Atta and Acromyrmex symbionts are highly and obligatorily polyploid (ca. 5–7 haplotypes on average). This stepwise transition appears analogous to ploidy variation in plants and fungi domesticated by humans and in fungi domesticated by termites and plants, where gene or genome duplications were typically associated with selection for higher productivity, but allopolyploid chimerism was incompatible with sexual reproduction.     

Minimizing the biodiversity impact of Neotropical oil palm development

Oil palm agriculture is rapidly expanding in the Neotropics, at the expense of a range of natural and seminatural habitats. A key question is how this expansion should be managed to reduce negative impacts on biodiversity. Focusing on the Llanos of Colombia, a mixed grassland–forest system identified as a priority zone for future oil palm development, we survey communities of ants, dung beetles, birds and herpetofauna occurring in oil palm plantations and the other principal form of agriculture in the region – improved cattle pasture – together with those of surrounding natural forests. We show that oil palm plantations have similar or higher species richness across all four taxonomic groups than improved pasture. For dung beetles, species richness in oil palm was equal to that of forest, whereas the other three taxa had highest species richness in forests. Hierarchical modelling of species occupancy probabilities indicated that oil palm plantations supported a higher proportion of species characteristic of forests than did cattle pastures. Across the bird community, occupancy probabilities within oil palm were positively influenced by increasing forest cover in a surrounding 250 m radius, whereas surrounding forest cover did not strongly influence the occurrence of other taxonomic groups in oil palm. Overall, our results suggest that the conversion of existing improved pastures to oil palm has limited negative impacts on biodiversity. As such, existing cattle pastures of the Colombian Llanos could offer a key opportunity to meet governmental targets for oil palm development without incurring significant biodiversity costs. Our results also highlight the value of preserving remnant forests within these agricultural landscapes, protecting high biodiversity and exporting avian ‘spill‐over’ effects into oil palm plantations.     

Comparative morpho-physiology of the metapleural glands of two Atta leaf-cutting ant queens nesting in clayish and organic soils

Queens of leaf-cutting ants found their nests singly, each consisting of a vertical tunnel and a final horizontal chamber. Because of the claustral mode of nest founding, the queen and/or her initial fungus garden are exposed to threats imposed by several soil pathogens, and the antibiotic secretions produced by their metapleural glands are considered a main adaptation to deal with them. Nests of two Atta leaf-cutting ant species, Atta vollenweideri and Atta sexdens rubropilosa, occur in different soil types, alfisols and oxisols. Their queens are known to excavate the initial nest in different soil horizons, clayish and organic, respectively, which differ in their fertility and associated microbiota. The aim of the present study was to comparatively investigate the morpho-physiology of the metapleural glands in queens of A. vollenweideri and A. sexdens rubropilosa, addressing the question whether the distinct selective pressure imposed by the microbiota in the two different soil types led to morpho-physiological differences in the metapleural glands that were consistent with their antiseptic function. The results revealed that metapleural glands of A. sexdens rubropilosa have a larger number of secretory cells, and consequently a higher production of antibiotic secretions, which may have been selected to allow nest founding at the superficial horizon of oxisols rich in organic matter and microorganisms. Glands of A. vollenweideri, on the contrary, presented fewer secretory cells, suggesting less production of antibiotic secretions. We argue that the excavation of deep founding nests in A. vollenweideri was primarily selected for during evolution to avoid the risk posed by flooding, and further hypothesize that a reduced number of cells in their metapleural glands occurred because of a weak pathogen-driven selective pressure at the preferred soil depth.     

Endemic predators, invasive prey and native diversity

Interactions between native diversity and invasive species can be more complex than is currently understood. Invasive ant species often substantially reduce diversity in the native ants diversity that act as natural control agents for pest insects. In Indonesia (on the island of Sulawesi), the third largest cacao producer worldwide, we show that a predatory endemic toad (Ingerophrynus celebensis) controls invasive ant (Anoplolepis gracilipes) abundance, and positively affects native ant diversity. We call this the invasive-naivety effect (an opposite of enemy release), whereby alien species may not harbour anti-predatory defences against a novel native predator. A positive effect of the toads on native ants may facilitate their predation on insect vectors of cacao diseases. Hence, toads may increase crop yield, but further research is needed on this aspect. Ironically, amphibians are globally the most threatened vertebrate class and are strongly impacted by the conversion of rainforest to cacao plantations in Sulawesi. It is, therefore, crucial to manage cacao plantations to maintain these endemic toads, as they may provide critical ecosystem services, such as invasion resistance and preservation of native insect diversity.