Larval recognition by Temnothorax longispinosus and T. ambiguus hosts of the slave-making ant Protomognathus americanus revisited: colony-level referent ensures conspecific preference

Slave-making ants exploit the labour of their own or another species. Temnothorax ambiguus and T. longispinosus are both ant species that serve as hosts of the obligatory slave-making ant Protomognathus americanus and are facultative slave-makers themselves. We offered laboratory colonies of T. ambiguus and T. longispinosus a series of choices among different larval types to better understand their brood discrimination abilities. Workers of both species preferentially accepted nestmate over non-nestmate larvae. Both species preferentially retrieved unrelated conspecific larvae over congeneric allospecific larvae, and T. ambiguus workers consumed more allospecifics than conspecifics. When presented with conspecific versus P. americanus larvae, both species manifested a clear bias towards conspecific larvae in terms of earlier retrieval and reduced cannibalism. That workers did not prefer P. americanus larvae over conspecific larvae as documented in previous research likely reflects the fact that in the present study, subject workers had access to the entirety of their colony as a referent during rearing and at the time of testing, as they would in nature. Moreover, reciprocal contact between P. americanus and conspecific larvae increased acceptance of the slave-maker larvae, but did not appear to lessen the acceptability of conspecific larvae. This suggests that transfer of cues through contact may be sufficient to alter the recognition signature of P. americanus larvae increasing acceptability by their hosts.     

Convergent evolution of the Dufour’s gland secretion as a propaganda substance in the slave-making ant genera Protomognathus and Harpagoxenus

Slave-making ants are social parasites which exploit the workforce of heterospecific slaves for their own reproduction, and to this end they have developed a variety of morphological and behavioural adaptations. Furthermore, social parasites utilize the chemical communication system of their hosts by breaking their nestmate recognition code, and some slave-maker species additionally employ semiochemicals as weapons during colony foundation and slaveraiding. Here, we demonstrate the use of such a ‘propaganda allomone’ by the North American myrmicine slave-maker Protomognathus americanus. This substance is produced in the Dufour’s gland and may be employed during slave raids to elicit panic among defending host workers. Slave-maker Dufour’s gland secretions evoked agitation and heightened levels of activity among host workers when applied directly on a host nest, and strong aggressive responses of nestmates when applied onto a host worker. Although the hosts own Dufour’s gland secretion also elicits intra-colonial fights, no support for the hypothesis was found that the slave-maker propaganda substance mimics a fertility signal of the host, as the chemical profile of the gland secretions is highly divergent between the two species. Preliminary results on the chemical composition of the secretion obtained by gas chromatography indicate that the propaganda substance of P. americanus differs from that of the related European slavemaker Harpagoxenus sublaevis, and is thus likely to represent an independent evolutionary development. Received 24 February 2005; revised 28 July 2005 and 2 March 2006; accepted 6 March 2006.     

Blochmannia endosymbionts improve colony growth and immune defence in the ant Camponotus fellah

Background  

Microorganisms are a large and diverse form of life. Many of them live in association with large multicellular organisms, developing symbiotic relations with the host and some have even evolved to form obligate endosymbiosis [1]. All Carpenter ants (genus Camponotus) studied hitherto harbour primary endosymbiotic bacteria of the Blochmannia genus. The role of these bacteria in ant nutrition has been demonstrated [2] but the omnivorous diet of these ants lead us to hypothesize that the bacteria might provide additional advantages to their host. In this study, we establish links between Blochmannia, growth of starting new colonies and the host immune response.     

Intracolonial conflict in the slave-making ant <Emphasis Type="Italic">Protomognathus americanus</Emphasis>: dominance hierarchies and individual reproductive success

Summary Social group viability results from a trade-off between cooperation and conflict, driven respectively by group and individual interests. Workers of the slave-making ants are known to have a high egg-laying potential, leading to a potential conflict over male production. Queenright and queenless nests of the slave-making ant Protomognathus americanus show a near-linear dominance hierarchy, and dominance rank is correlated with reproductive activity. Genetic and behavioural analysis revealed that the queen, when present in the nest, is behaviourally dominant and monopolises reproduction. In queenless nests, the haploid (male) brood is produced primarily by a single worker. We suggest the dominance hierarchy regulates male production, between the queen and her workers as well as among workers. Comparison of our results to another study allows us to place our data in an ecological context. This slave-making ant species appears to fit the concession model of reproductive skew: where resources (i.e. host nests) are poor, there is strong skew and where resources are richer reproduction is more egalitarian.Received 31 July 2003; revised 7 October 2003; accepted 9 October 2003.     

Habitat-related microgeographic variation of worker size and colony size in the ant Cataglyphis cursor

In social insects, colony size is a crucial life-history trait thought to have major implications for the evolution of social complexity, especially in relation to worker size polymorphism. Yet, little is known about how ecological factors can affect and constrain colony. Here, we explored the pattern of colony-size and worker-size variation in the Mediterranean ant Cataglyphis cursor, in relation to the type of habitats colonized (seaside vs. vineyard). The high level of the water table in the seaside habitat could constrain the depth of C. cursor underground nests and directly constrain its colony size. If worker size increases with colony size, as observed in other ant species, larger colony size and larger workers should be found in the vineyard populations. By comparing worker size among 16 populations, we verified that workers were significantly larger in the vineyard populations. We further determined that the morphological similarities detected among populations from the same habitat type were not due to geographic or genetic proximity. In two populations from each habitat type, the depth of nests was positively correlated with colony size and colony size with worker size. Using a type II regression approach, we further showed that the difference between the two populations in the depth of nest was sufficient to explain the difference in colony size, and similarly, variation in colony size was sufficient to explain variation in worker size. Our results suggest that a single proximate ecological factor could lead to significant variation in major life-history parameters.     

Cuticular profiles and mating preference in a slave-making ant

The remarkable ability of slave-making ants to integrate chemically in the colonies of their host species makes them useful model systems for investigating the role of cuticular hydrocarbons in chemical recognition. The purpose of our study was to examine the influence of the rearing host species on the cuticular hydrocarbon profile and on the mating behaviour of sexuals of the slave-making ant Chalepoxenus muellerianus. Sexuals from a population parasitizing exclusively the host species Temnothorax unifasciatus were reared in the laboratory either with their natural host or another potential host species, Temnothorax recedens. C. muellerianus males reared with T. unifasciatus investigated and mounted female sexuals reared with the same host significantly more often than female sexuals reared with T. recedens. Similarly, C. muellerianus males reared with T. unifasciatus discriminated against female sexuals from natural T. recedens colonies. Males experimentally or naturally reared with T. recedens did not clearly discriminate between female sexuals reared by the two host species and only rarely engaged in mating attempts with either type of female sexuals. Chemical analyses showed that host species affect the chemical profile of C. muellerianus sexuals and vice versa. Our findings indicate that cuticular hydrocarbons might be important in the mating success of this ant species. Received 21 June 2006; revised 20 February 2007; accepted 1 March 2007.     

Parasite prevalence and the size of host populations: an experimental test

Although important in epidemiological theory, the relationship between the size of host populations and the prevalence of parasites has not been investigated empirically. Commonly used models suggest no relationship, but this prediction is sensitive to assumptions about parasite transmission. In laboratory populations, I manipulated the size of Tribolium castaneum flour beetle populations and measured the prevalence and distribution of a parasitic mite, Acarophenax tribolii. I found that parasite prevalence did not vary for a wide range of host population sizes. However, prevalence was lower in populations with less than 40 hosts. This effect cannot be attributed to changes in host population density because host density was held constant among treatments. The reduction in prevalence of small populations below a threshold that I observed is predicted by the extinction debt model, but it is not expected from models of host-parasite interactions that assume density-dependent transmission. The distribution of parasites, measured using Lloyd's patchiness index, was not affected by host population size. The mean crowding of parasites, however, was negatively related with host density. Finally, the prevalence of parasites in large populations did not differ from that found in sets of smaller patches as long as the smaller populations in aggregate were equivalent in size to the large population.     

Body size but not colony size increases with altitude in the holarctic ant,Leptothorax acervorum

1. Bergmann's rule states that organisms inhabiting colder environments show an increase in body size or mass in comparison to their conspecifics living in warmer climates. Although originally proposed for homoeothermic vertebrates, this rule was later extended to ectotherms. In social insects, only a few studies have tested this rule and the results were ambiguous. Here, ‘body size’ can be considered at two different levels (the size of the individual workers or the size of the colony). 2. In this study, data from 53 nests collected along altitudinal gradients in the Alps were used to test the hypotheses that the worker body size and colony size of the ant Leptothorax acervorum increase with increasing altitude and therefore follow Bergmann's rule. 3. The results show that the body size of workers but not the colony size increases with altitude. Whether this pattern is driven by starvation resistance or other mechanisms remains to be investigated.     

Quantifying the effect of colony size and food distribution on harvester ant foraging

Desert seed-harvester ants, genus Pogonomyrmex, are central place foragers that search for resources collectively. We quantify how seed harvesters exploit the spatial distribution of seeds to improve their rate of seed collection. We find that foraging rates are significantly influenced by the clumpiness of experimental seed baits. Colonies collected seeds from larger piles faster than randomly distributed seeds. We developed a method to compare foraging rates on clumped versus random seeds across three Pogonomyrmex species that differ substantially in forager population size. The increase in foraging rate when food was clumped in larger piles was indistinguishable across the three species, suggesting that species with larger colonies are no better than species with smaller colonies at collecting clumped seeds. These findings contradict the theoretical expectation that larger groups are more efficient at exploiting clumped resources, thus contributing to our understanding of the importance of the spatial distribution of food sources and colony size for communication and organization in social insects.     

Aggressive behaviours of female Drosophila melanogaster are influenced by their social experience and food resources

Abstract Aggressive behaviours of female Drosophila melanogaster (Meigen) (Diptera: Drosophilidae) were studied in the laboratory. These behaviours included 'approach', a seemingly intentional movement of a fly to another fly, 'lunge', a dash at another fly, and 'wings erect', a brief erection of wings toward another fly. These behaviours were influenced by social experience and food resources. First, event frequencies of aggressive behaviours were higher in flies that had been held in isolation than in those that had been held in crowded conditions. This difference was the result of the social experience of attacking flies and not that of attacked flies. Second, flies showed aggressive behaviours more frequently when they were on food containing yeast colonies than when they were on food without yeast colonies. Third, flies conditioned to a food solution containing live yeast prior to observation showed lower frequency of aggressive behaviours than flies conditioned to a boiled food solution. Development of aggressive behaviours became apparent one day after eclosion.     

Cuticular spectra in the slave-making ant Polyergus rufescens and the slave species Formica rufibarbis

Abstract. The relative proportions of cuticular components having the same retention times were compared between the slave-making ant Polyergus rufescens and the slave ant Formica rufibarbis living in monospecific or mixed colonies. The two species were found to present different spectra. The Formica workers, when enslaved by Polyergus , tend to lose their colony characteristics but they do not seem to adopt the characteristics of Polyergus.     

Recent changes in colony sizes of two seabird species in Scotland are not strongly influenced by Birds Directive status of the colony

Special Protection Areas (SPAs) designated under the EU Birds Directive have improved the conservation status of many terrestrial bird species in Europe, but protecting breeding sites may be less effective for highly mobile birds such as seabirds. Colony census data for Great Skuas Stercorarius skua and Great Black-backed Gulls Larus marinus in Scotland show that breeding numbers have fared no better in sites where these species are SPA breeding features and, counter-intuitively, the evidence indicates better performance in non-SPA colonies, most likely because non-SPA colonies are generally smaller so are less subject to density-dependent competition. The main drivers of population change are widespread rather than colony-based in these two species with recent reductions in carrying capacity. Many other seabird species are vulnerable to similar widespread pressures so seabird conservation strategy needs to focus on mitigating these pressures, as designation of seabird breeding sites as SPAs is not enough to ensure effective seabird conservation.     

Invasive alien plants affect grassland ant communities, colony size and foraging behaviour

Ants are dominant members of many terrestrial ecosystems and are regarded as indicators of environmental changes. However, little is known about the effects of invasive alien plants on ant populations, particularly as regards the density, spatial distribution and size of ant colonies, as well as their foraging behaviour. We addressed these questions in a study of grassland ant communities on five grasslands invaded by alien goldenrods (Solidago sp.) and on five non-invaded grasslands without this plant. In each grassland, seven 100 m² plots were selected and the ant colonies counted. Ant species richness and colony density was lower in the plots on the invaded grasslands. Moreover, both of these traits were higher in the plots near the grassland edge and with a higher number of plant species in the grasslands invaded by goldenrods but not in the non-invaded ones. On average, ant colony size was lower on the invaded grasslands than the non-invaded ones. Also, ant workers travelled for longer distances to collect food items in the invaded areas than they did in the non-invaded ones, even after the experimental removal of some ant colonies in order to exclude the effect of higher colony density in the latter. Our results indicate that invasive alien goldenrods have a profound negative effect on grassland ant communities which may lead to a cascade effect on the whole grassland ecosystem through modification of the interactions among species. The invasion diminishes a major index of the fitness of ants, which is a colony’s size, and probably leads to increased foraging effort of workers. This, in turn, may have important consequences for the division of labour and reproductive strategies within ant colonies.     

City limits: Heat tolerance is influenced by body size and hydration state in an urban ant community

Cities are rapidly expanding, and global warming is intensified in urban environments due to the urban heat island effect. Therefore, urban animals may be particularly susceptible to warming associated with ongoing climate change. We used a comparative and manipulative approach to test three related hypotheses about the determinants of heat tolerance or critical thermal maximum (CT_max) in urban ants—specifically, that (a) body size, (b) hydration status, and (c) chosen microenvironments influence CT_max. We further tested a fourth hypothesis that native species are particularly physiologically vulnerable in urban environments. We manipulated water access and determined CT_max for 11 species common to cities in California's Central Valley that exhibit nearly 300‐fold variation in body size. There was a moderate phylogenetic signal influencing CT_max, and inter (but not intra) specific variation in body size influenced CT_max where larger species had higher CT_max. The sensitivity of ants’ CT_max to water availability exhibited species‐specific thresholds where short‐term water limitation (8 hr) reduced CT_max and body water content in some species while longer‐term water limitation (32 hr) was required to reduce these traits in other species. However, CT_max was not related to the temperatures chosen by ants during activity. Further, we found support for our fourth hypothesis because CT_max and estimates of thermal safety margin in native species were more sensitive to water availability relative to non‐native species. In sum, we provide evidence of links between heat tolerance and water availability, which will become critically important in an increasingly warm, dry, and urbanized world that others have shown may be selecting for smaller (not larger) body size.     

Selective acceptance of the brood of two formicine slave-making ants by host and non-host related species

Summary In a laboratory choice-test, free-living ant workers ofFormica cunicularia andF. rufibarbis (subgenusServiformica), both potentially slave species of the obligatory slave-makerPolyergus rufescens, cared for cocoons of this parasite and for homocolonial cocoons at comparable rates. Both potential hosts did not differ in their capacity to rear the parasite brood. This fact is discussed in relation to host selection and specificity inP. rufescens. No such attraction and/or tolerance was found towards cocoons of the facultative slave-makerFormica sanguinea, which also enslaves both host species. Workers ofF. lugubris, a species which is never enslaved, destroyed cocoons from both slave-making species. The attractiveness of the brood ofP. rufescens for both potentially slave species could be due to an interspecific brood pheromone in addition to brood mimicry. An alternative hypothesis is a close phylogenetic distance between this slave-maker andServiformica species. The capacity to gain acceptance by adult slave workers might be one of the crucial evolutionary steps separating obligatory from facultative slave-making ants.     

Effects of group size and pine defence chemicals on Diprionid sawfly survival against ant predation

The defence chemicals and behavioural adaptations (gregariousness and active defensive behaviour) of pine sawfly larvae may be effective against ant predation. However, previous studies have tested their defences against very few species of ants, and few experiments have explored ant predation in nature. We studied how larval group size (groups of 5 and 20 in Neodiprion sertifer and 10, 20 and 40 in Diprion pini) and variation in levels of defence chemicals in the host tree (Scots pine, Pinus sylvestris) affect the survival of sawfly larvae. Food preference experiments showed that ants do eat sawfly larvae, although they are not their most preferred food item. According to our results, ant predation significantly increases the mortality rate of sawfly larvae. Larval mortality was minor on pine tree branches where ant traffic was excluded. We also found that a high resin acid concentration in the host tree significantly decreased the mortality of D. pini larvae when ants were present. However, there was no such relationship between the chemical concentrations of the host tree and larval mortality for N. sertifer. Surprisingly, grouping did not help sawfly larvae against ant predation. Mortality risk was the same for all group sizes. The results of the study seemingly contradict previous understanding of the effectiveness of defence mechanisms of pine sawfly against ant predation, and suggest that ants (Formica exsecta in particular) are effective predators of sawfly larvae.An erratum to this article can be found at