Queen adoption in the polygynous and polydomous ant, Leptothorax curvispinosus

Alate female reproductives of the facultatively polygynous andpolydomous ant, Lepiothorax curuispinosus, were reared fromfield-collected nests, mated, and introduced into either theirparental nests or alien conspecific nests. The 41 queens introducedinto alien nests were usually attacked and rejected (97. 6%),but one queen was accepted after initial aggression. The 27queens introduced into their parental nests received a variableresponse. Some were accepted without any apparent aggression(59. 3%), but others were strongly attacked and rejected (40.7%). Sequential introductions of up to four queens into particularparental nests indicated that nests consistently either acceptor reject their mated offspring. The presence or absence ofresident queens in parental nests had no apparent influenceon the acceptance of offspring queens. Nests that accepted queenshad significantly fewer workers than those that did not, butthis slight difference is unlikely to explain these dichotomousresults and could be spurious. Dissections of the introducedqueens revealed that 79. 0% were inseminated and 98. 3% haddeveloping, yolked eggs in their ovarioles, but these variableshad no apparent effect on acceptability. Similarly, the sizeof the introduced queens and the time that elapsed between matingand introduction had no apparent effect. The consistent responseof parental nests in either accepting or rejecting their matedoffspring indicates a mechanism of queen number regulation inthis species that involves characteristics of the colony ornest rather than variability among offspring queens. This mechanismcould be responsible for maintaining relatively low numbersof queens and high genetic relatedness in colonies (or individualnests) while promoting flexibility in colony reproduction bycolony fission ("budding") and the dispersal of young queens.This mechanism could also involve an important conflict of interestbetween parental colonies and their mated offspring and mightcontribute to the evolution of socially parasitic colony foundationstrategies. Acceptance of mated offspring by their parentalcolonies might only occur during certain periods in colony development,depend on the level of genetic diversity within the colony (ornest), reflect the condition of the colony, nest, queen(s),brood, or local habitat, or result from a genetic polymorphism.     

Both female castes contribute to colony emigration in the polygynous ant Mystrium oberthueri

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Queen execution in a monogynous ant

Workers in many species of social insects are capable of laying unfertilized eggs, which can develop into haploid males. This causes a conflict about male parentage between queens and workers. In a few species, this may result in matricide, that is, workers kill the colony''s queen. Queen killing has so far been observed mainly in multi‐queen colonies or in annual species, when the queen''s fecundity declines at the end of the reproductive period. Here, we report queen expulsion and matricide in a monogynous, monandrous ant with perennial societies. Workers were seen to aggressively expel both related and unrelated queens from their nest shortly after the end of hibernation. Queen expulsion and matricide led to a significant decrease in the number of workers and brood, but eventually increased the direct fitness of workers through significant male production. Long‐term observations revealed a short lifespan of queens, while workers in orphaned colonies survived and produced male offspring over several years.     

Nest site limitation and colony takeover in the ant Leptothorax nylanderi

Ecological constraints on the success of independent coloniesare thought to strongly shape the organization of ant societies.One of the most important factors is probably the availabilityof suitable empty nest sites. By population censuses, laboratoryexperiments, and microsatellite analyses, we investigated thecolony and population structure of the small, myrmicine antLeptothorax (Myrafant) nylanderi in a deciduous forest nearWürzburg, Germany, where nest sites appear to be stronglylimited, especially in late summer. Colonies of L. nylanderiinhabit cavities in rotting branches, hollow acorns, grass stems,etc. After hibernation, a temporary overabundance of empty nestsites facilitates the fragmentation of larger colonies intosmaller buds, which, because the species is monogynous, arein part queenless. Nest sites become scarce in summer due torapid deca and both established colonies and young foundingqueens face a severe shortage of suitable nest sites. This leadsto the fusion of established, unrelated colonies, which afterinitial fighting permanently merge and live together. Typicalyonly one queen survives after fusion. Similarly, young matedqueens may seek adoption in alien nests instead of foundingtheir own colonies solitarily, and here again only a singlequeen survives. This temporary intraspecific parasitism maybe an important first step in the evolution of obligatory permanentparasitism, which is widespread in the genus Leptothorax.     

The economics of brood raiding and nest consolidation during ant colony founding

Summary The most dangerous time for an ant colony is during the founding stage when the small colony is vulnerable to predation and competition. Colonies can grow more rapidly when multiple queens cooperate in raising the first worker brood (pleometrosis) or by raiding other incipient colonies for their brood. This brood raiding has been proposed to be the primary force selecting for pleometrosis, i.e. multiple-queen colonies may have a considerable advantage in destroying neighbours by aggressively stealing their brood. An alternative hypothesis is that incipient nests are part of a larger, interconnected population structure and that brood raiding reflects cooperative pleometrosis with subdivided colonies. A simple mathematical model supports the second hypothesis: workers of incipient colonies are especially favoured to peaceably abandon their nest and join with other colonies if the queens are related or queens from raided colonies can infiltrate the raiding colony. The latter condition is often met in ant species that brood raid and particularly exemplified in fire ants (Solenopsis invicta), where brood raiding involves little mortal combat and combines with pleometrosis to rapidly increase colony size. It is proposed that the term nest consolidation should replace brood raiding to more accurately reflect the relatively non-aggressive and potentially apparently cooperative nature of interactions between incipient ant colonies.     

Queen recruitment in a multiple-queen population of the fire ant Solenopsis invicta

We assessed patterns of new queen recruitment in a polygyne(multiple queens per nest) population of the fire ant Solenopsisinvicta in its introduced range. Newly recruited queens wereidentified using four physiological markers, and genotypic datafrom nuclear and mitochondrial markers were used to estimaterelatedness of new nest mate queens to each other and to theolder nest mate queens. In total, 1.7% of the queens collectedin late spring and early summer were deemed to be new recruits.The relatedness of these queens to each other and to the olderqueens within nests was not significantly different from zero,suggesting that newly recruited queens represent a random sampleof potential reproductive queens in the population. Moreover,the number of new queens recruited within nests was not correlatedwith the number of older queens present and did not differ significantlyfrom a Poisson distribution. Thus, queen recruitment in this populationof S. invicta appears to occur at random with respect to thenumber of older queens present within nests.     

Sex allocation in a facultatively polygynous ant: between-population and between-colony variation

We investigated sex allocation in three U.K. populations ofthe facultatively polygynous ant Leptothorax acervorum over1-3 years. The first main finding was that, across sites, thepopulation sex-investment ratio changed from significantly femalebiased to significantly male biased with increasing polygyny.This was consistent with workers controlling sex allocationand reacting to changes in their population-level relatedness asymmetry.It was also consistent with local resource competition due to reproductionby colony budding under polygyny. Worker control was supportedby the finding that queen number had no effect on sex allocationamong polygynous colonies. The second main result was that monogynouscolonies consistently produced more female-biased sex-investmentratios than polygynous colonies in one site only (Santon). Theresults from Santon supported both the relative relatednessasymmetry hypothesis and the idea of sex ratio compensationdue to colony budding. The workers' response to their population-levelrelatedness asymmetry reinforced the case for relatedness asymmetrybeing influential at the colony level. The other populationscould have lacked split sex ratios because polygynous colonieswere either comparatively rare or common, making them behaveas almost entirely monogynous (Aberfoyle) or polygynous (Roydon) populations.In Roydon, this was consistent with the inference from allozyme datathat monogynous and polygynous colonies did not differ in theirworker relatedness asymmetries. The final principal findingwas that, of hypotheses linking the colony sex-investment ratiowith sexual productivity, there was support for the constantfemale hypothesis but not for the constant male, cost variation,or multifaceted parental investment hypotheses.     

The underdog invader: Breeding system and colony genetic structure of the dark rover ant (Brachymyrmex patagonicus Mayr)

Ants are among the most successful species at invading new environments. Their success undeniably comes from their various modes of reproduction and colony breeding structures, which influence their dispersal ability, reproductive potential, and foraging strategies. Almost all invasive ant species studied so far form supercolonies, a dense network of interconnected nests comprising numerous queens, without aggression toward non‐nestmates. This strategy results in invasive colonies that are able to grow extremely fast and large while avoiding intraspecific competition, allowing them to monopolize environmental resources and outcompete native species. Here, we developed and used 10 microsatellite markers to investigate the population structure and breeding system of the dark rover ant Brachymyrmex patagonicus Mayr in its introduced range. We determined whether this species exhibits a supercolonial structure by assessing whether different nests belonged to the same genetic colony. We inferred its dispersal ability by investigating isolation by distance and estimated the numbers of queens per colonies and mating per queen through parent‐offspring inferences. We found that most of the colonies of B. patagonicus were comprised of a single nest, headed by a single queen. Each nest was distinct from one another, without isolation by distance, which suggests strong dispersal ability through nuptial flights. These features are commonly observed in noninvasive and native ant species, but they are surprising for a successful invasive ant, as they strongly differ from other invasive ants. Overall, we discuss how this seemingly unfavorable strategy for an invasive ant might favor the invasive success of the dark rover ant in the United States.     

Self-organized asymmetries in ant foraging: a functional response to food type and colony needs

The dominant paradigm to explain asymmetries in the spatialdistribution of foraging animals is that they track the spatialheterogeneity of their environment. However, in social insects,endogenous spatial asymmetries can emerge within a uniformenvironment as an outcome from the self-organizing processof trail recruitment. We studied how self-organized asymmetries contribute to the exploitation of different food sources (carbohydrateor proteins) in colonies of the aphid-tending ant Lasius nigervarying in their nutritional needs (presence or absence ofbrood). Colonies with brood fed on sucrose sources exhibita higher mobilization of foragers than the other experimentalgroups. Foraging patterns differ greatly according to food type: colonies strongly focus their activity on only one dropletof sucrose, whereas they show a rather homogeneous distributionof foragers between proteinaceous sources. In addition, thepresence of brood in the colony enhances the asymmetry of collectiveforaging for both types of food. These spatial differencesin self-organized foraging patterns allow efficient exploitationof natural resources and play a role in the competitive strategy of this widespread palearctic ant.     

Predatory behavior of a seed-eating ant:Brachyponera senaarensis

The great flexibility of the feeding strategies exhibited by the ponerine ant Brachyponera senaarensis (Mayr) allows it to exploit either seeds or animal prey items as food resources. Predation is generally limited to small prey and is very similar to scavenging behavior. In laboratory conditions, the predatory behavior of B. senaarensis is not different in structure from that known in other carnivorous ants species. The workers forage individually and return to the nest using a series of cues involving light, a chemical graduated marking system near the nest entrance, and memory. During nest-moving, recruitment by tandem running was observed. However, in colonies where the food supply is regular, workers that discover food do not recruit nestmates, but make repeated trips between the nest and the food source. On the contrary, in starved colonies, the introduction of prey may produce a massive exit of foragers, corresponding to a primitive form of mass recruitment similar to that observed in some other ant species.     

Brood development into sexual females depends on the presence of a queen but not on temperature in an ant dispersing by colony fission,Aphaenogaster senilis

Abstract. 1. In eusocial insects, colony fission is a mode of dispersal by which a young queen leaves her nest with some workers to found a new colony. In these species, adult females (workers and the queen) should allocate most resources to increasing their colony size, which constrains the possibility of fission. In contrast, developing diploid larvae should have a preference for becoming a queen and having their own offspring, rather than becoming workers and rearing the offspring of other females. 2. In the ant Aphaenogaster senilis, queens are produced in very small numbers, suggesting that adult females control larval development. We used a 6‐year series of data on more than 300 nests to determine the annual cycle of worker and queen production. Although both overlapped, the latter mostly occurred in the second half of the summer, after a major peak of worker emergence. Young queens were also often produced in nests whose reproductive queen had died, thus allowing her replacement. Overall, we estimate that only 0.07% of diploid larvae actually develop into gynes. 3. Laboratory experiments indicated that brood is bipotent until the second larval instar. Diploid larval development into queen was favoured by the removal of the mother queen, but was not affected by rearing temperature. 4. Our data suggest that most diploid broods are forced by the adults to develop into workers rather than into gynes. However, when the queen is not present due to death or after a fission event, a few larvae are allowed to develop into gynes. One way for workers to limit the development of larvae might be by controlling the amount of food they receive.     

Six weeks in the life of a reproducing army ant colony: male parentage and colony behaviour

The army ant Eciton burchellii is one of the most conspicuous ant species in New World tropical forests, but studies of colony life histories have been hampered by the nomadic lifestyle of these ants, which alternate between a nomadic phase when the colony relocates frequently, and a statary phase when the colony remains at a fixed site. Here we report on a colony from Venezuela that we studied continuously for six weeks, from the time that the queen produced a reproductive brood until the adult reproductives emerged and the colony entered the next cycle. Our findings support the contention that reproductive larvae develop faster than worker larvae, and that the nomadic phases of colonies with reproductive broods are significantly shorter than those of colonies with worker broods. This strongly suggests that the onset of pupation is linked to the onset of the statary phase. We used microsatellite genotyping to accurately identify male and queen larvae and we describe how they can be distinguished morphologically. Using the same genetic markers, we determined the parentage of 81 males produced by this colony. Only one of the males had a genotype that could not be directly derived from the observed queen genotype, but this mismatch is most probably due to a single mutation at one of the microsatellite loci, rather than this male being a worker son. We therefore conclude that this colony provides no evidence that workers lay eggs that develop into adult males in the presence of the queen, confirming the results of an earlier study on male parentage in an Old World army ant. Received 16 November 2006; revised 15 January 2007; accepted 16 January 2007.     

Changing colony growth rates inCamponotus floridanus as a behavioral response to conspecific presence (Hymenoptera: Formicidae)

Previous work with the antCamponotus floridanus demonstrated that perception of competition can be clearly differentiated from effects of mortality and decreased resources. That is, brood biomass in ant colonies decreases as a consequence of a behavioral decision(s) rather than because of limited food availability or reduced numbers of brood tenders. The experiments presented here extend that work. Under experimental conditions, colony growth inC. floridanus is modified by distance between brood and unrelated conspecifics and by worker age distribution. When nonnestmates are encountered at the nest versus at a separate foraging site, less brood is maintained by a colony. Although colonies with older workers maintain a brood biomass similar to that of colonies with younger workers, that biomass is concentrated in fewer, larger, more rapidly maturing larvae. These effects seem to be due entirely to worker control.     

Population recovery and spatial dynamics of colony recruitment in a harvester ant: a case study in a park

The recovery process of Messor aciculatus (Fr. Smith) harvester ant colonies from habitat disturbance by construction work in an urban park was investigated from 1989 to 1994. Colonies located in 1989 were considered to be survivors of the disturbance, and colonies located for the first time after 1990 were considered to be recruits. The colonies continuously increased in number after the disturbance, and the population size more than doubled just after the disturbance. Analyses of the nest site distribution of colonies in 1989 and 1994 suggest that landform modification destroyed some of them, and that the changed nest site distribution in 1989 affected the distribution in 1994, or, in other words, the disturbance indirectly affected the nest distribution 5 years later. Analyses of the nearest neighbor to each colony suggested that established colonies would obstruct newly recruited colonies around their nests. Construction may open up space for recruitment colonization by removing established colonies, and may also create suitable conditions by destruction of original vegetation, or returning vegetation to the early stages of plant community succession. As a result, the disturbance may facilitate the subsequent recruitment of new colonies and rejuvenate the population.     

Worker demography and behavior in a supercolonial ant colony: The case of the desert ant Cataglyphis niger

Insect societies headed by multiple queens (polygyny) raise evolutionary questions, such as how does genetic heterogeneity among colony members affect in-nest interactions; or, are all queens equally reproductive or equally treated by workers? Answering such questions requires intensive and continuous observations of in-nest behavior. Here, we addressed these questions in the polygyne supercolonial ant, Cataglyphis niger, using a barcoding system that enables the tracking of individual interactions, together with polymorphic DNA microsatellite markers that indicate the matriline and patriline of all individuals. Our findings that both queens and workers have low interrelatedness corroborate earlier reports regarding the supercolony structure of C. niger. Ovary inspection and worker genotyping revealed that all the queens contribute similarly to nest demography. Tracking positions of individual workers through time revealed that only a small proportion of them are constantly engaged in tending the queens and can be considered as retinue workers. However, genotyping these workers and the attended queens revealed no relationship to genetic relatedness, again typical of a true polygyne and supercolonial species. Unlike invasive supercolonial species, C. niger is native to Israel, enabling us to address questions regarding the driving forces, other than kin selection, that stabilize this society.     

Effects of nest size and dispersion on brood production in a North American population of wood ant Formica fusca (Hymenoptera: Formicidae)

We examined several key parameters of the population ecology of a North American population of Formica fusca (L.), including nest dispersion, colony size and brood production. Physical nest size was significantly correlated with colony size, and colony size, in turn, was significantly correlated with brood production. Sex allocation was male biased, although larger nests were more likely to produce reproductive female brood (gynes). Neither nest temperature nor moisture level was significantly correlated with brood production. Formica fusca nests in this population had a comparatively low average nearest-neighbor distance with a significantly even pattern of dispersion, which suggests relatively high intraspecific competition. However, nearest-neighbor distance was not significantly associated with either colony size or relative brood production.     

Specialized predation on plataspid heteropterans in a coccinellid beetle: adaptive behavior and responses of prey attended or not by ants

Two plataspid hemipteran species proliferated on Bridelia micrantha(Euphorbiaceae). Colonies of Libyaspis sp., never attendedby ants, developed on branches, while Caternaultiella rugosa lived at the base of the trunks, mostly in association withCamponotus brutus that attends them in carton shelters. Bothplataspid species are prey of the coccinellid beetle Anisolemniatetrasticta, whose larvae always detected them by contact.When attacked the Libyaspis nymphs cowered, so that the hypertrophiedlateral sides of their tergits made contact with the substrate,but the ladybirds slid their long forelegs under these nymphs,lifted them, and bit them on the ventral face. The Caternaultiellanymphs, which do not have hypertrophied extremities of thetergits, tried to escape at contact with the ladybirds, butwere rarely successful. To capture them, the ladybirds eitheradopted the previous behavior or directly grasped then bitthem. We noted a graded aggressiveness in the ants toward theladybirds according to the situation: no aggressiveness onthe tree branches; stopping the ladybirds that approached theshelters where the ants attended Caternaultiella; and fullattack of ladybirds that tried to capture Caternaultiella nymphssituated outside shelters. The latter behavior can emit analarm pheromone that triggers the dispersion of their congenerswhile attracting attending C. brutus workers. Naive workersare not attracted, so we deduce that this behavior is the resultof a kind of learning.     

Reproductive conflict in social insects: male production by workers in a slave-making ant

In insect societies, workers cooperate but may also pursue their individual interests, such as laying viable male eggs. The case of obligatory slave-making ants is of particular interest because workers do not engage in maintenance activities and foraging. Therefore, worker egg laying is expected to be less detrimental for colony efficiency than in related, nonparasitic species. Furthermore, as slave-making workers usually do not perform brood care and thus might have little power in manipulating sex allocation, they might be more strongly selected to increase their direct fitness by producing their own sons than workers in nonparasitic species. In this study we investigated worker reproduction in four natural colonies of the slave-making ant Polyergus rufescens, using highly variable microsatellite markers. Our results show that workers produce up to 100% of the males. This study thus presents the first direct evidence of an almost complete takeover of male reproduction by workers in ants.     

The army ant Aenictus silvestrii and its related species in Southeast Asia,with a description of a new species (Hymenoptera: Formicidae: Aenictinae)

Three species of Aenictus with reduced numbers of antennal segments (8–9) occur in Southeast Asia: A. jarujini sp. nov., A. latifemoratus and A. silvestrii. Aenictus latifemoratus is new to Java and Borneo, while A. silvestrii is recorded for the first time from Borneo.     

No sympatric speciation here: multiple data sources show that the ant Myrmica microrubra is not a separate species but an alternate reproductive morph of Myrmica rubra

No aspect of speciation is as controversial as the view that new species can evolve sympatrically, among populations in close physical contact. Social parasitism has been suggested to yield necessary disruptive selection for sympatric speciation. Recently, mitochondrial DNA phylogeography has shown that the ant Myrmica microrubra is closely related to its host, Myrmica rubra, leading to the suggestion that sympatric speciation has occurred. We investigated the relationships between the two ant forms using mitochondrial and nuclear DNA sequences, microsatellite genotyping and morphometrics. Molecular phylogenetic and population structure analyses showed that M. microrubra does not evolve separately to its host but rather shares a gene pool with it. Probability analysis showed that mitochondrial DNA data previously adduced in favour of sympatric speciation do not in fact do so. Morphometrically, M. microrubra is most readily interpreted as a miniature queen form of M. rubra, not a separate species. Myrmica microrubra is not an example of speciation. The large (typical M. rubra) and small (M. microrubra) queen forms are alternative reproductive strategies of the same species. Myrmica microrubraSeifert 1993 is consequently synonymized here with M. rubra Linnaeus, 1758.