Ant attendance in aphids: why different degrees of myrmecophily?

1. Aphids show a range of associations with ants from nonattendance to obligate myrmecophily. Aphis fabae cirsiiacanthoides is facultatively associated with ants, while Symydobius oblongus is an obligate myrmecophile. The selection pressures that have shaped these associations are unknown. 2. The consequences for these aphids of their different degrees of associations with ants were determined, in terms of costs and benefits to individuals and colonies in laboratory and field experiments. In the laboratory, individuals of A. f. cirsiiacanthoides performed worse and those of Symydobius oblongus performed better when attended by the ant Lasius niger than when unattended. For example, when ant-attended, A. f. cirsiiacanthoides developed more slowly, were smaller, and invested less in gonads, whereas S. oblongus developed more quickly, were larger, and invested more in gonads. In addition, the ant regulated the population size of S. oblongus to an average of 50–70 individuals per birch sapling by removing aphids, but did not regulate the population size of A. f. cirsiiacanthoides. 3. Under field conditions, ant-attended colonies of both A. f. cirsiiacanthoides and S. oblongus achieved higher peak numbers and lasted longer, and ant-attended colonies of A. f. cirsiiacanthoides produced more alate dispersers than unattended colonies. 4. The implications of divergent selection pressures for the development of myrmecophily in aphids are discussed.     

Genome‐wide transcriptome signatures of ant‐farmed Squamellaria epiphytes reveal key functions in a unique symbiosis

Farming of fungi by ants, termites, or beetles has led to ecologically successful societies fueled by industrial‐scale food production. Another type of obligate insect agriculture in Fiji involves the symbiosis between the ant Philidris nagasau and epiphytes in the genus Squamellaria (Rubiaceae) that the ants fertilize, defend, harvest, and depend on for nesting. All farmed Squamellaria form tubers (domatia) with preformed entrance holes and complex cavity networks occupied by P. nagasau. The inner surface of the domatia consists of smooth‐surfaced walls where the ants nest and rear their brood, and warty‐surfaced walls where they fertilize their crop by defecation. Here, we use RNA sequencing to identify gene expression patterns associated with the smooth versus warty wall types. Since wall differentiation occurred in the most recent common ancestor of all farmed species of Squamellaria, our study also identifies genetic pathways co‐opted following the emergence of agriculture. Warty‐surfaced walls show many upregulated genes linked to auxin transport, root development, and nitrogen transport consistent with their root‐like function; their defense‐related genes are also upregulated, probably to protect these permeable areas from pathogen entry. In smooth‐surfaced walls, genes functioning in suberin and wax biosynthesis are upregulated, contributing to the formation of an impermeable ant‐nesting area in the domatium. This study throws light on a number of functional characteristics of plant farming by ants and illustrates the power of genomic studies of symbiosis.     

Testing the optimal defense hypothesis in Stryphnodendron adstringens (Fabaceae,Mimosoideae) leaves: the role of structure,number, position and nectar composition of extrafloral nectaries

Stryphnodendron adstringens is a common Cerrado tree that possesses extrafloral nectaries (EFNs) on its leaves, which are located at the base and apex of the rachis and along the secondary veins. The position of EFNs and their nectar production can be affected by defense strategies because plant organs possess different values and herbivory vulnerability. Here we aimed to elucidate anatomy, histochemistry, nectar composition and EFN number on leaves of S. adstringens in the light of the optimal defense hypothesis. We found a convergence on anatomy and histochemical characterization because the three studied types of EFNs have epidermis, secretory parenchyma and vascular tissue, showing phenolic compounds and polysaccharides in the secretory parenchyma cells. The nectar contained glucose, fructose and sucrose, which attract ants of the Camponotus and Cephalotes genus. We found differences in the number of EFNs along the secondary veins and in the nectar composition between EFNs located at the base and apex of the rachis of the leaf. The number of EFNs on the secondary veins increases from the base to the apex, suggesting a strategy to induce ant patrolling over the entire leaf region. EFNs at the base secreted more nectar, which should be related to the protection of the leaf base, which is the part most vulnerable to herbivore attack and the most valued organ. We concluded that EFNs of S. adstringens are anti‐herbivore defenses whose pattern matches the predictions of the optimal defense hypothesis.     

Bacteria may contribute to distant species recognition in ant–aphid mutualistic relationships

Mutualistic interactions between ant and aphid species have been the subject of considerable historical and contemporary investigations, the primary benefits being cleaning and protection for the aphids and carbohydrate‐rich honeydew for the ants. Questions remained, however, as to the volatile semiochemical factor influencing this relationship. A recent study highlighted the role of bacterial honeydew volatile compounds in ant attraction. Here, ant's ability to distantly discriminate 2 aphid species was investigated based on bacterial honeydew semiochemicals emissions using a two‐way olfactometer. Both the mutualistic aphid Aphis fabae L. and the nonmyrmecophilous aphid Acyrthosiphon pisum Harris were found to be attractive for the ant Lasius niger L. The level of attraction was similar in both assays (control vs. one of the aphid species). However, when given a choice between these 2 aphid species, ants showed a significant preference for Aphis fabae. Honeydew volatiles, mostly from bacterial origins, are known to be a key element in ant attraction. Using the same olfactometry protocol, the relative attractiveness of volatiles emitted by honeydews collected from each aphid species and by bacteria isolated from each honeydew was investigated. Again, ants significantly preferred volatiles released by Aphis fabae honeydew and bacteria. This information suggests that microbial honeydew volatiles enable ants to distantly discriminate aphid species. These results strengthen the interest of studying the occurrence and potential impact of microorganisms in insect symbioses.     

Five polymorphic microsatellite markers for the study of Cardiocondyla elegans (Hymenoptera: Myrmicinae)

We describe primer sequences for five microsatellite markers in Cardiocondyla elegans, an ant species with ergatoid males. Polymorphism of these loci was investigated using 236 individuals from 22 colonies from four locations. The microsatellites are dinucleotide repeats with four to 16 alleles, and the observed heterozygosity ranges from 0.244 to 0.720. We characterized these markers for the study of the population as well as the social structure of colonies.     

Impacts of Argentine ants on avian nesting success

Biological invasions can have severe and widespread impacts on ecological communities. A few species of ants have become particularly damaging invaders but quantitative data of their impacts on many taxa is still lacking. We provide experimental evidence using artificial nests baited with quail eggs that the invasive Argentine ant (Linepithema humile) can be a significant avian nest predator – Argentine ants recruited to more nests and in higher abundance than the native ant species they displace. However, at a site invaded by Argentine ants, we monitored over 400 nests of a ground-nesting species, the Dark-eyed Junco (Junco hyemalis), and found that less than 2% of nests failed as a result of Argentine ant predation/infestation. A review of the literature also suggests that Argentine ants may not be a serious threat to bird nests relative to other predators or parasites. However, invasive ants with the capability of overwhelming prey though stinging (specifically the red-imported fire ant, Solenopsis invicta), may have a higher impact on avian nesting success. Received 14 January 2005; revised 28 April 2005; accepted 12 May 2005.     

Genetic relationships among populations of the Argentine ant Linepithema humile introduced into Japan

Since some colonies of Argentine ant Linepithema humile were discovered in Japan in 1993, populations of this invasive alien ant species have been expanding their distribution. To resolve the number of invasions and the genetic structure in the early stages of introduction, we inferred the genetic structure and relationships among colonies from eight localities, from analyses of eight nuclear microsatellite DNA markers. F _ST analysis, principal component analysis and assignment test showed that at least three highly genetically differentiated groups of Argentine ants are present in Japan. Populations from Hiroshima Bay were grouped together (Hiroshima, Hatsukaichi, Otake, Iwakuni Central and Iwakuni Kuroiso), while those from Kobe and Yanai were both genetically distant from each other and from the Hiroshima Bay group. Hatsukaichi and Kobe are international seaports, suggesting that the by-ship invasion occurred at least twice. The invasion route of the Yanai population is unknown at this moment. The Aichi population was genetically distant from that of the Hiroshima Bay group by the difference in allele frequencies, and it was plausible that the Aichi population was introduced from the Hiroshima Bay group by human-mediated jump dispersal.     

Isolation of polymorphic microsatellite loci in the ponerine ant Ectatomma tuberculatum

We report the development of 11 microsatellite markers by an enrichment protocol in the ponerine ant Ectatomma tuberculatum. Polymorphism was explored in two colonies collected from Mexico and two colonies from Brazil. From the 11 loci which amplified, seven loci showed intracolonial polymorphism in Mexican colonies and only six loci were variable in populations from Brazil. Observed heterozygosity ranged from 0.18 to 0.84. The other five loci exhibited different alleles between Mexico and Brazil but geographical variability was not investigated further. Cross‐amplification was tested in another species of the same Tribe (Gnamptogenys striatula) and one locus was revealed to be polymorphic.     

Immediate impacts of invasion by Wasmannia auropunctata (Hymenoptera: Formicidae) on native litter ant fauna in a New Caledonian rainforest

Abstract For the last 30 years, Wasmannia auropunctata (the little fire ant) has spread throughout the Pacific and represents a severe threat to fragile island habitats. This invader has often been described as a disturbance specialist. Here we present data on its spread in a dense native rainforest in New Caledonia. We monitored by pitfall trapping the litter ant fauna along an invasive gradient from the edge to the inner forest in July 1999 and March 2000. When W. auropunctata was present, the abundance and richness of native ants drops dramatically. In invaded plots, W. auropunctata represented more than 92% of all trapped ant fauna. Among the 23 native species described, only four cryptic species survived. Wasmannia auropunctata appears to be a highly competitive ant that dominates the litter by eliminating native ants. Mechanisms involved in this invasive success may include predation as well as competitive interactions (exploitation and interference). The invasive success of W. auropunctata is similar to that of other tramp ants and reinforces the idea of common evolutionary traits leading to higher competitiveness in a new environment.     

Floral volatiles controlling ant behaviour

1 . Ants show complex interactions with plants, both facultative and mutualistic, ranging from grazers through seed predators and dispersers to herders of some herbivores and guards against others. But ants are rarely pollinators, and their visits to flowers may be detrimental to plant fitness.
2 . Plants therefore have various strategies to control ant distributions, and restrict them to foliage rather than flowers. These 'filters' may involve physical barriers on or around flowers, or 'decoys and bribes' sited on the foliage (usually extrafloral nectaries - EFNs). Alternatively, volatile organic compounds (VOCs) are used as signals to control ant behaviour, attracting ants to leaves and/or deterring them from functional flowers. Some of the past evidence that flowers repel ants by VOCs has been equivocal and we describe the shortcomings of some experimental approaches, which involve behavioural tests in artificial conditions.
3 . We review our previous study of myrmecophytic acacias, which used in situ experiments to show that volatiles derived from pollen can specifically and transiently deter ants during dehiscence, the effects being stronger in ant-guarded species and more effective on resident ants, both in African and Neotropical species. In these plants, repellence involves at least some volatiles that are known components of ant alarm pheromones, but are not repellent to beneficial bee visitors.
4 . We also present new evidence of ant repellence by VOCs in temperate flowers, which is usually pollen-based and active on common European ants. We use these data to indicate that across a wide range of plants there is an apparent trade-off in ant-controlling filter strategies between the use of defensive floral volatiles and the alternatives of decoying EFNs or physical barriers.