High Divergence of Cuticular Hydrocarbons and Hybridization Success in Two Allopatric Seven-Spot Ladybugs.

The seven-spotted ladybug is a widespread species in the Palearctic, and also acclimated in the Nearctic. It has been classified into different species on the basis of certain morphological characteristics, the geographical origin, and the genitalia structure of both sexes. The morphotypes of North Africa and the Canary Islands are separated, under the name of Coccinella algerica Kovár, 1977, from the rest of the Palearctic and Nearctic populations of Coccinella septempunctata Linnaeus, 1758. In this study, we investigated, on one hand, whether potential reproductive barriers have been established during evolution between the geographically isolated North African and the European seven-spotted ladybugs by performing reciprocal crosses. On the other hand, we assessed their cuticular hydrocarbon (CHC) divergence by GC-MS. The 33 CHCs indentified are with a skeleton of 23 to 32 carbon atoms. These CHCs are linear alkanes (24.9 ± 3.6%) and methyl-branched alkanes (75.1 ± 3.6%) including monomethylalkanes (48.8 ± 2.4%), dimethylalkanes (24.6 ± 4.0%) and trimethylalkanes (2.0 ± 1.0%). Although all the CHC compounds identified are present in the two seven-spotted ladybugs and their F1 and F2 hybrids, their profiles diverged significantly. However, these chemical divergences have not altered the sexual communication to cause reproductive isolation. The two ladybugs interbreed and leave viable and fertile offspring, with even a heterosis effect on reproductive performances, without phenotypic degradation after the F1 generation. So, these chemical differences are just an intraspecific variability in response to heterogeneous environments. The two types of ladybugs can be considered as two different races of the same species with reduced genetic divergence.

Effect of the land area elevation on the collective choice in ants.

Collective decisions regarding food source exploitation in social insects are influenced by a range of parameters, from source quality to individual preference and social information sharing. Those regarding the elevation of the physical trail towards a food source have been neglected. In this work, we investigated the effect of ascending and descending paths from the nest to a food source on collective choice in two ant species Lasius niger and Myrmica rubra. Our hypothesis that returning loaded with food from the high source is more energy efficient was validated by choice experiments: when the sources are simultaneously introduced the high food source is preferentially exploited by both species. The flexibility of colony response was then tested by introducing the preferred source (high) incidentally, after recruitment towards the down food source began. Despite the well-known lack of flexibility of L. niger, both species showed the ability to reallocate their foraging workforce towards the highest food source. The collective choice and the flexibility are based on the difference between the u-turn rates when foragers are facing the ascending or descending branch. We discuss these results in terms of species-specifics characteristics and ecological context.

Dispersal strategies in the highly polygynous ant Crematogaster (Orthocrema) pygmaea Forel (Formicidae: Myrmicinae).

In ants, dispersal strategies and morphology of female sexuals are generally linked to the mode of colony founding. In species using long-range dispersal tactics, queen/worker dimorphism is generally high and young queens are able to initiate new colonies by themselves, using their metabolic reserves. By contrast, in species using short-range dispersal strategies, queen/worker dimorphism is generally low and, due to their limited metabolic reserves, queens have lost the capacity to raise their brood alone and to found their colony independently. Moreover, polygyny is also often associated with short-range dispersal strategies, although the relationship between the number of queens and the dispersal strategy in ants is not clear-cut. Here, dispersal strategies were investigated in C. pygmaea, a highly polygynous and polydomous ant species from northeastern Brazil. Field observations and laboratory experiments show that this ant exhibits a suite of traits that are more commonly associated with long-range dispersal and independent colony foundation: functional wings in both males and females, high queen/worker dimorphism, strong weight loss in mature queens, nuptial flights and, in the lab, ability of young queens to found new colonies in haplometrotic conditions. On the other hand, this species shows a high degree of polygyny with a strong seasonal component, and, at least under laboratory conditions, mature queens seem able to develop propagules if they are accompanied by at least 10 workers. These features strongly suggest that (1) some of the gynes do not engage in a long-range dispersal but become new queens in their mother colony and (2) that budding events are possible in this species. We therefore speculate that C. pygmaea has a dual dispersal strategy probably related to environmental conditions: some gynes engage in long-range dispersal followed by independent colony foundation at the beginning of rainy season, while others mate in the parental colony and are re-adopted leading to high polygyny. During the rainy season, budding events can lead to colony extension and increased polydomy. Polydomy is commonly thought to improve resource discovery and exploitation through decentralized foraging behavior, a significant advantage during the rainy season when food ressources (mainly floral/extrafloral nectaries and hemipteran honeydew) are more abundant and when colony needs for food supplies are highest.

Social Structure and Genetic Distance Mediate Nestmate Recognition and Aggressiveness in the Facultative Polygynous Ant Pheidole pallidula.

In social insects, the evolutionary stability of cooperation depends on the privileged relationships between individuals of the social group, which is facilitated by the recognition of relatives. Nestmate recognition is based on genetically determined cues and/or environmentally derived chemical components present on the cuticle of individuals. Here, we studied nestmate recognition in the ant Pheidole pallidula, a species where both single-queen (monogyne) and multiple-queen (polygyne) colonies co-occur in the same population. We combined geographical, genetic and chemical analyses to disentangle the factors influencing the level of intraspecific aggressiveness. We show that encounters between workers from neighbouring colonies (i.e., nests less than 5 m away) are on average less aggressive than those between workers from more distant colonies. Aggressive behaviour is associated with the level of genetic difference: workers from monogyne colonies are more aggressive than workers from polygyne colonies, and the intensity of aggressiveness is positively associated with the genetic distance between colonies. Since the genetic distance is correlated with the spatial distance between pairs of colonies, the lower level of aggression toward neighbours may result from their higher relatedness. In contrast, the analysis of overall cuticular hydrocarbon profiles shows that aggressive behaviour is associated neither with the chemical diversity of colonies, nor with the chemical distances between them. When considering methyl-branched alkanes only, however, chemical distances differed between monogyne and polygyne colonies and were significantly associated with aggressiveness. Altogether, these results show that the social structure of colonies and the genetic distances between colonies are two major factors influencing the intensity of agonistic behaviours in the ant P. pallidula.

Queens defense by workers in the highly polygynous ant Crematogaster pygmaea (Hymenoptera: Myrmicinae).

Some aspects of the biology of Crematogaster pygmaea, a highly polydomous and polygynous ant, are more commonly found in monogynous species. One such characteristic is the high attractiveness of its queens. In this study, this attractiveness was assessed under varying experimental conditions to investigate the factors responsible for its expression and variation, and to identify the nature of queen attractiveness. It was shown (1) that C. pygmaea queens are highly attractive to the workers that cluster on and around them (retinue), (2) that the attractiveness of C. pygmaea queens is context-dependent, i.e., it increases with increasing degree of potential danger to the queen, (3) that the attractiveness signal of C. pygmaea queens is chemically based, and (4) that this signal is persistent and apparently not colony-specific. The proposed hypothesis is that the C. pygmaea queens constantly release an attractiveness signal that is "read" by the workers, in a dependent way linked to the context, and that the main function of this attractiveness is to protect queens. This protection would have a high adaptive value in the context of the social structure and the reproductive strategies in C. pygmaea.

The differential response of workers and queens of the ant Lasius niger to an environment marked by workers: ants dislike the unknown.

It is well known that ants can use cuticular hydrocarbons (CHCs) as a specific recognition cue. Most previous studies addressed the perception of CHCs occurring on the cuticle. However, the presence of CHCs in the environment (e.g., on the substrate) and the role of these compounds as a signal cue are less clear. In this study of the ant L. niger, CHCs were extracted from corpses of workers, from the legs of workers and from filter paper marked by workers. Behavioural choice tests were then performed to study the reactions of workers and queens to CHC passively deposited on the substrate by walking workers. A GC-MS analysis of the CHCs revealed a complex mixture of 28 peaks composed of linear alkanes, alkenes and mono-, di- or trimethylalkane. This mixture was consistently present in the three sources tested (corpses of workers, the tarsus of workers and paper marked by walking workers). Nevertheless, discriminant analyses distinguished clearly among the three types of profiles. The results of our behavioural experiments showed that the workers were able to distinguish between marked and unmarked papers but that they exhibited no preferences for different types of marked papers (papers marked by workers from their own colony or papers marked by workers from a different colony). In queens, the perception of marked papers depended on age or physiological state. Just-mated queens did not discriminate between unmarked papers and papers marked by an alien colony. In contrast, the egg-laying queens 3 days after mating significantly favoured the marked papers. We discuss the ecological relevance of the differences in decision-making between castes.

Genetics, behaviour and chemical recognition of the invading ant Pheidole megacephala.

Introduced species often become ecologically dominant, displacing native species and posing a serious threat to ecosystem function and global biodiversity. Ants are among the most widespread and damaging alien species; introductions are often accompanied by population-level behavioural and genetic changes contributing to their success. We investigated the genetic structure, chemical profile and nestmate recognition in introduced populations of the invasive big-headed ant, Pheidole megacephala, in Australia. Behavioural analyses show that workers are not aggressive towards conspecifics from different nests, even at large geographical scales (up to 3000 km) and between populations encompassing a wide range of environmental conditions. By contrast, interactions with workers of other species invariably result in agonistic behaviours. Genetic analyses reveal that populations have low genetic diversity. No genetic differentiation occurs among nests of the same population; differentiation between populations, though significant, remains weak. Chemical analyses indicate that cuticular lipids are similar between colonies of a population, and that differentiation between populations is low. Altogether, these results indicate that the big-headed ant P. megacephala forms a large unicolonial population across northern/eastern Australia.

Ovarian activity correlates with extreme changes in cuticular hydrocarbon profile in the highly polygynous ant, Linepithema humile.

In social insects, cuticular hydrocarbons of adults may vary with ovarian activity. Such variations are suggested to function in the regulation of reproduction within colonies. The modification of the CH profile with ovarian activity is usually interpreted as a signal of fertility causing workers to refrain from reproducing in the presence of the queen. We examined the effect of ovarian activity on the CH profiles in the Argentine ant Linepithema humile, a species where workers lack ovaries and are completely sterile. Our data show considerable differences in the CH profiles between fertile and infertile individuals. These differences are mainly qualitative. The CH profile strongly changes at the start of egg laying, both in mated and unmated queens. These results show that variation in cuticular hydrocarbons with ovarian activity in ants is not restricted to species with worker reproduction. We propose that in the Argentine ant, the cuticular hydrocarbons of laying queens correspond to a signal of fertility involved in the regulation of various aspects of reproduction, such as the rearing of new sexuals.