Nest Founding Research Articles

Prehibernating aggregations of Polistes dominulus: an occasion to study early dominance assessment in social insects

At a very early age several mammals establish a first dominance hierarchy, which often persists into adulthood. Social wasps offer an excellent opportunity to study such a phenomenon in insects. Indeed, foundresses of several paper wasps meet in clusters to hibernate from September to March. In spring, wasps found new associative nests where linear hierarchies occur. In the first phase of hibernation, clustering Polistes dominulus wasps show most of the social interactions occurring on the nest 6 months later. At the emergence from diapause, some females already show some behavioral and physiological traits typical of dominant individuals. Here, we investigated the significance of the interactions in the autumnal clusters. We demonstrated that in a given pair, it is more likely that the dominant wasp in autumn becomes the alpha female in spring after the nest foundation phase occurred. Moreover, we showed that dominant females in clusters have both larger body size and ovaries. As ovarian development mainly depends on the social context, our findings seem to indicate that social factors affect the tendency to dominate in aggregations. Furthermore, we suggest that some females may reinforce their physiological status by dominating in clusters, thus increasing the probability to become dominant in spring.

Mating flight, metrosis, and semi-claustrality in the seed-harvester ant Pogonomyrmex salinus (Hymenoptera, Formicidae)

Here we examine dispersal, metrosis, and claustrality in the seed-harvester ant Pogonomyrmex salinus at an unusually large mating aggregation. We found that mode of queen dispersal from the mating aggregation is not a function of queen mass and that wing damage among queens was relatively rare. P. salinus is haplometrotic in the field and foundress queens placed together in forced associations eventually fight to the death. While queens of Pogonomyrmex salinus can survive claustrally, producing a single minim from their body reserves in the laboratory, fed queens produce up to four significantly larger minims along with concurrent larvae and pupae during the same period. Since queens forage in the field, we interpret claustrality as a secondary reserve strategy when foraging fails, and suggest that foraging is obligate for P. salinus queens in an overdispersed and temperate environment. Thus, nest founding strategies employed by P. salinus may be environmentally determined and represent a continuum between fully claustral and obligate foraging. We discuss our results with reference to theories of pleometrosis and claustral colony founding.

It's good to be queen: classically eusocial colony structure and low worker fitness in an obligately social sweat bee

Lasioglossum malachurum, a bee species common across much of Europe, is obligately eusocial across its range but exhibits clear geographic variation in demography and social behaviour. This variation suggests that social interactions between queens and workers, opportunities for worker oviposition, and patterns of relatedness among nest mates may vary considerably, both within and among regions. In this study, we used three microsatellite loci with 12-18 alleles each to examine the sociogenetic structure of colonies from a population at Agios Nikolaos Monemvasias in southern Greece. These analyses reveal that the majority of colonies exhibit classical eusocial colony structure in which a single queen mated to a single male monopolizes oviposition. Nevertheless, we also detect low rates of multiqueen nest founding, occasional caste switching by worker-destined females, and worker oviposition of both gyne and male-producing eggs in the final brood. Previous evidence that the majority of workers show some ovarian development and a minority (17%) have at least one large oocyte contrasts with the observation that only 2-3% of gynes and males (the so-called reproductive brood) are produced by workers. An evaluation of the parameters of Hamilton's Rule suggests that queens benefit greatly from the help provided by workers but that workers achieve greater fitness by provisioning and laying their own eggs rather than by tending to the queen's eggs. This conflict of interest between the queen and her workers suggests that the discrepancy between potential and achieved worker oviposition is due to queen interference. Comparison of relatedness and maternity patterns in the Agios Nikolaos Monemvasias population with those from a northern population near Tübingen, Germany, points to a north-south cline of increasingly effective queen control of worker behaviour.

Production of Minima Workers by Gynes of Atta colombica Guérin-Ménéville (Hymenoptera: Formicidae: Attini) that Lack a Fungal Pellet

Abstract Early studies of Atta nest founding showed that gynes exhibit claustral nest foundation, cultivate fungal gardens with fecal secretions, and nourish their larvae with eggs. These studies also showed that gynes sometimes lose their fungal pellets, or the fungal garden fails before workers emerge, apparently dooming the incipient colony. Here we report that Atta colombica foundresses maintained in the laboratory can produce workers even though they lack fungal pellets to initiate gardens. If such behavior occurs in nature, it raises the possibility that workers might re-acquire a fungal symbiont after nest establishment, and potentially rescue failing colonies.

Restricted gene flow between two social forms in the ant Formica truncorum

We studied genetic differentiation between two social forms (M-type: single queen, independent nest founding; P-type: multiple queens, dependent nest founding, building of colonial networks) of the ant Formica truncorum in a locality where the social types characterize two sympatric populations. The genetic results indicate restricted gene flow between the social forms. Female gene flow between the forms appears to be absent as they did not share mitochondrial haplotypes. Significant nuclear differentiation and the distribution of private alleles suggest that male gene flow between the forms is weak. However, the assignment analysis indicates some gene flow with P males mating with M females. The results have potentially important implications concerning social evolution within the forms but they need to be confirmed in other localities before they can be generalized. The colonies in the M-type population have earlier been shown to produce split sex ratios, depending on the mating frequency of the queens. The inferred gene flow from the P to the M type means that the split sex ratio is partly suboptimal, possibly because the P populations are not long-lived enough to influence the behavioural decisions in the M colonies.

Facultative polygyny in Ectatomma tuberculatum (Formicidae, Ectatomminae)

Polygyny, the presence of several mated queens within the same colony, is widespread in insect societies. This phenomenon is commonly associated with ecological constraints such as limited nest sites. In habitats where solitary nest foundation is risky, monogynous colonies can reintegrate young daughter queens (secondary polygyny). We studied the reproductive structure (i.e. queen number) of the ectatommine ant Ectatomma tuberculatum from Bahia State, Brazil. This species was found to present facultative polygyny: out of a total of 130 colonies collected, 39.2% were monogynous, while 43.8% were polygynous. Polygynous colonies had significantly more workers than monogynous ones. Queen number in polygynous colonies ranged from 2 to 26, with an average of 4 ± 4 queens per colony. All nestmate queens were egg-layers with no apparent dominance hierarchy or agonistic behavior. Non-nestmate queens were adopted by monogynous colonies suggesting that polygyny is secondary, originating through queen adoption. This species is characterized by an open recognition system, which probably allows a switch from monogynous to polygynous colonies. The behavioral acts of queens showed that resident queens remained frequently immobile on or near the brood, contrarily to alien or adopted queens and gynes. In addition, monogynous queens showed no behavioral or physiological (i.e. by ovarian status) differences in comparison with polygynous ones. Secondary or facultative polygyny, probably associated with queen adoption, may have been favored in particular environmental conditions. Indeed, by increasing colony productivity (i.e. number of workers) and territory size (by budding and polydomy), polygyny could uphold E. tuberculatum as a dominant species in the mosaic of arboreal ants in Neotropical habitats.

Demography of the Okinawan eusocial waspRopalidia fasciata(Hymenoptera: Vespidae) I. Survival rate of individuals and colonies, and yearly fluctuations in colony density

AbstractWe conducted a series of demographic studies of the primitively eusocial waspRopalidia fasciatain Okinawa, a subtropical part of Japan, and found the following. The colony cycle of this wasp is annual, spanning from April to November or even December; this is longer than the colony cycles of other temperate polistine wasps so far reported. The survival rate of the marked foundresses was 40% to June, dropping to 1.4% by September. These survival rates are similar to other subtropical and tropical species. Most females that emerged in November were probably second generation adults (progeny of the original foundresses), which would participate in founding nests in the following spring. These facts indicate thatR. fasciatain Okinawa is at least partially bivoltine. Survival of a nest to September was 10–20%; however, because a failed nest is often rebuilt, survival of the colony to September was as high as 50%. The mean number of new foundresses produced per foundress was 7.5, and their overwintering survival was 16%. Hence, a single foundress produced, on average, 1.2 progeny foundresses to the following year. Density dependence was shown in the rate at which the progeny foundresses were produced. These results explain the remarkable stability of nest densities from year to year in the area. The above results reveal thatR. fasciatain Okinawa shares many demographic characteristics with other primitively eusocial wasps, particularly year‐to‐year stability of nest density and a long colony cycle.

Queen phenotype and behaviour during cooperative colony founding in Polyrhachis moesta

Founding queens of the arboreal ant Polyrhachis moesta aggregate independently of kinship and cooperate in caring for their brood. In field studies, the number of queens in a founding nest varied from 1 to 8. The number of queens in the nests with multiple queens decreased significantly with time after the nuptial flight, resulting in monogynous or oligogynous nests. Single- and multiple-founding queens did not differ in characteristics representing nutritional states or body size immediately after the nuptial flight. Wet body weight decreased as days passed, whereas head width of founding queens who overwintered successfully were relatively larger. In laboratory studies, founding queens performed liquid food exchanges more frequently with queens from other founding nests or immature colonies than with those from the same nests. Queens in founding nests and immature colonies were observed to show no aggression against non-nestmate queens, whereas queens in established colonies showed aggressive behaviours against non-nestmates. This indicates that founding queens change drastically in their aggression levels before and after colony establishment. Multiple-founding queens started laying eggs earlier than single-founding queens under laboratory conditions. Higher brood productivity and lower brood mortality were observed in multiple-queen nests. These potential advantages in multiple-queen founding may support the cooperative association among unrelated founding queens.

Solitary nesting and reproductive success in the paper wasp Polistes aurifer

Female Polistes paper wasps are capable of independent nesting, yet many populations demonstrate a mixture of solitary and cooperative nest foundation. Previous studies of Polistes have found survival and/or productivity advantages of cooperative nest foundation compared to solitary nesting, and reproductive skew models have been designed to predict the dynamics of such flexible cooperation. In this paper, we examine the success of different nesting strategies in a previously unstudied population of Polistes aurifer in southern California. The colony cycle of this population is less synchronous than that of other temperate species, and the frequency of solitary nesting averages 86.2%. Our data suggest that this low rate of cooperative nest founding is adaptive, as demonstrated by the lack of survival or productivity advantages for cooperative foundress associations. Due to foundress turnover and nest foundation later in the season, many nests produce only one set of offspring. This results in a loss of the eusocial nature of some nests in the population. Data from a small sample of multifoundress nests show significant positive reproductive skew, despite concession model predictions that skew should be low in populations with low ecological constraints on independent nesting. This lack of support for the concessions skew model reflects a diminished incentive for cooperation.

Hibernation clustering and its consequences for associative nest foundation in Polistes dominulus (Hymenoptera Vespidae)

Various studies indicate that cuticular hydrocarbons (CHCs) act as cues for nestmate recognition in paper wasps and other social insects. A recent study showed that associative nest foundation in Polistes dominulus is mainly performed by foundresses coming from the same locality. In the present study, we induced future foundresses of P. dominulus collected in two different localities to hibernate in the laboratory in aggregates of individuals from the same or different localities. After 2 months of hibernation, foundresses from different localities but from the same experimental cluster did not show any preference to associate, at the time of nest foundation, with individuals coming from the same original locality. The cuticular chemical profiles of individuals from the mixed hibernation clusters were quite similar and significantly different from those of individuals which hibernated with other foundresses from the same locality. These findings suggest that, in this species, mechanisms other than nestmate chemical recognition play a major role in the spring association of gynes during the foundation of a new nest.

Development of harvester ant colonies alters soil chemistry

Nests of the harvester ant Pogonomyrmex barbatus typically contain higher concentrations of organic matter, nitrogen and phosphorus than surrounding soils. The difference between nest soils and surrounding soils is due, at least in part, to ant foraging behavior. Ants retrieve seeds from the environment and concentrate seed nutrients in the vicinity of the nest. But elevated nutrient concentrations in nests may also reflect initial conditions, if nest-founding queens are more likely to choose or survive in soils with high organic matter and nutrient content. By measuring the soil nutrient content and surface area of P. barbatus nests ranging from 1 to 20 years of age over two sampling periods, we (a) investigated the relationship between nest soil characteristics and colony age, and (b) tested the hypothesis that nest soils differ from background soils when nests are established. Nest surface area increased with colony age until age 5–10 years and leveled off thereafter. Relative to surrounding soils, concentrations of total nitrogen and orthophosphate increased, and pH decreased, with increasing colony age. The difference between nest soils and surrounding soils in total nitrogen, nitrate, and ammonium concentrations also increased over a 9-month interval between sampling bouts. Extrapolations from regressions of soil chemical variables against colony age provided no evidence that nest founding and early colony survival is more likely to occur in high-nutrient soils.

Ecological traits and evolutionary sequence of nest establishment in fungus-growing ants (Hymenoptera, Formicidae, Attini)

Neotropical ants in the tribe Attini share the obligatory behaviour of cultivating fungi as an essential food source. Of this complex and well-studied mutualism, little is known about the biological traits of the phylogenetically basal attines, which may offer important clues to the origins and early evolution of this intricate symbiosis. In this paper we focus on the founding of new nests, a key to understanding evolutionary changes in many social insects, which has received comparatively little attention in attine ants. We present a comparative survey of nest-founding behaviour in Attini, based on 441 foundress nests of 20 species in eight attine genera. In general, attine queens are semiclaustral and haplometrotic when founding nests. We show that attine foundresses of most species use an inert platform (discarded forewings, roots, or rocks) on which the incipient fungal garden is physically isolated during nest foundation, and they conduct semiclaustral foundation. This behaviour is not shared with Atta, which places the incipient garden directly on the soil floor of the initial nest chamber, and conducts claustral foundation. Nest-founding maps congruently onto the major clades in attine phylogeny, suggesting that the behavioural mechanisms used to isolate the incipient garden may have been key innovations in the early evolution of attine fungiculture. The evolutionary sequence of gardening in Attini suggests a transformation series from retention and use of detached wings (suspended from the chamber ceiling or placed over the chamber floor) to root-suspension, and finally use of bare soil. We also discuss transitions from semiclaustral to claustral founding, as well as from haplometrosis to pleometrosis, from ecological and evolutionary perspectives. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81, 39–48.

Impacto da queima controlada da cana-de-açúcar na nidificação e estabelecimento de colônias de Atta bisphaerica Forel (Hymenoptera: Formicidae)

Queimadas de baixas intensidades podem promover sensíveis alterações nas propriedades físicas, químicas e microbiológicas do solo, na profundidade onde se estabelecem os sauveiros iniciais de Atta Fabricius. Considerando que o estabelecimento da nova colônia é o período mais crucial da vida da colônia, investigaram-se neste trabalho as implicações pós-fogo de uma queima controlada de palhiço de cana-de-açúcar de intensidade igual a 615 kJ. m-1.s-1 na escolha de locais para escavação dos ninhos e estabelecimento inicial de colônias de Atta bisphaerica Forel. A queima eliminou matéria orgânica superficial da área, o que implicou em aumento significativo de densidade até a profundidade de 15 cm. Da mesma forma, a eliminação da vegetação pelo fogo promoveu maior resistência do solo à penetração. Essas alterações nas propriedades físicas não afetaram significativamente a escavação dos ninhos iniciais por fêmeas de A. bisphaerica e nem tampouco influenciaram a profundidade de construção de suas câmaras iniciais em condições de laboratório. A queima alterou a constituição química do solo e promoveu alterações na atividade microbiológica do mesmo. Em condições de campo, as fêmeas de A. biphaerica apresentaram maior taxa de estabelecimento em locais queimados. Em condições de laboratório, colônias cujas fêmeas estabeleceram-se em solos provenientes de áreas queimadas apresentaram maior mortalidade do que aquelas estabelecidas em solos de áreas não queimadas.

Variable worker behaviour in the weakly eusocial sweat bee, Halictus sexcinctus Fabricius

Studies of eusocial halictines suggest that workers have many reproductive options, including sterile altruism in the maternal nest, combined helping and personal reproduction, and diapause and spring nest founding. How and when workers exercise these various options influences the strength of colony social organization. Halictus sexcinctus exhibits highly polymorphic social behaviour, with solitary colonies in central Europe and both eusocial and communal colonies in southern Greece. Indirect evidence suggests that some worker-brood females are actually gynes. A distinctly bimodal size distribution among foundresses in 1998, the lower size peak being close to the modal body size of workers from 1997, suggests that large worker-brood females overwinter and return to the aggregation as eusocial foundresses. Other first-brood females remain in the maternal nest as workers, although few can be classified as classical, sterile altruists. Only 17% of older, healthy workers are sterile (i.e. had ovarian development scores ≤ 0.1), whereas about 83% are reproductive, exhibiting at least one 1/4-developed oocyte. About 57% of older workers have at least one fully or 3/4 developed oocyte, signifying that they are ready or almost ready to lay. Sterile workers exhibit greater total wear (combined mandibular and wing wear) scores than reproductive workers, suggesting that they are older, have higher activity rates, or both.

INITIAL NEST DEVELOPMENT IN THE SWARM-FOUNDING PAPER WASP, POLYBIA PAULISTA (HYMENOPTERA: VESPIDAE, EPIPONINI): CASES OF BUILDING MULTIPLE INITIAL COMBS

Early steps of nest foundation and development were studied in the Neotropical swarm-founding paper wasp, Polybia paulista. During early colony foundation P. paulista wasps gather into tight clusters adjacent to the newly established nest-site. While the period ranging from swarm settlement to nest construction took about 12 days in summer swarms, this period tended to be longer in winter swarms. In all colonies we observed, eggs were observed in at least one comb, which was not yet completely covered by the envelope. Among seven studied cases, two swarms built multiple initial combs, of which combs were fused into a single one within a few days. On account of rarity in Polybia, it's difficult to conclude if this unusual habit is anomalous, or not. However, since comb fusion develops so efficiently and rapidly, it seems likely to be included in the building repertory of the species under concern.

Benefits of foundress associations in the paper wasp Polistes dominulus: increased productivity and survival, but no assurance of fitness returns

Successful Polistes dominulus nests can be started by one or more nest founding queens (foundresses). Consequently, there is much interest in the specific benefits that induce cooperation among foundresses. Here, we experimentally demonstrate one major benefit of cooperation, namely that multiple foundresses increase colony productivity. This increase is close to the value predicted by subtracting the productivity of undisturbed single-foundress colonies from the productivity of undisturbed multiple-foundress colonies. However, we found no evidence that an associating foundress' contribution to colony growth is preserved if she disappears (assured fitness returns). Our correlational data suggest that cooperation provides survival benefits, multiple-foundress colonies are more likely to survive to produce offspring than are single-foundress colonies, and individual foundresses in multiple-foundress groups are less likely to disappear before worker emergence than foundresses nesting alone. Therefore, association provides substantial productivity and survival benefits for cooperating foundresses.

Cuticular hydrocarbons and reproductive status in the social wasp Polistes dominulus

. Most species of social insect are characterized by a reproductive division of labor among morphologically specialized individuals. In contrast, there exist many species where all individuals are morphologically identical and dominance relationships determine which individuals mate and/or reproduce. In newly founded multiple-foundress associations of the social wasp Polistes dominulus, foundresses establish dominance hierarchies where the top-ranked (alpha) female monopolizes egg laying. The possibility that chemical cues are used for recognition of egg-laying individuals has not been explored in this wasp. Using non-destructive techniques, we examined the relationship between ovarian activity and the proportions of cuticular hydrocarbons of three female types (dominant and subordinate foundresses and workers) in 11 colonies. Immediately after nest foundation, no differences were found between alpha and subordinate females. In contrast, at worker emergence, alpha females were statistically distinguishable from both subordinates and workers. We experimentally removed the alpha female in 5 of the original nests and reanalyzed hydrocarbon proportions of the new dominant individual. Replacement individuals were all found to acquire a cuticular signature characteristic of the alpha female. This suggests that cuticular hydrocarbons are used as cues of ovarian activity in P. dominulus, and we discuss our results in terms of a switch from behavioral dominance to chemical signaling in this wasp.

Kin Associations during Nest Founding in an Allodapine Bee Exoneura robusta: do Females Distinguish between Relatives and Familiar Nestmates?

True recognition of kin can have important fitness consequences in terms of directing altruistic behaviours toward close relatives (nepotism) and avoiding inbreeding. However, recent evidence suggests that some social insect species cannot or do not distinguish their closest relatives from among nestmates in important fitness-based contexts. Such findings are relevant to kin selection theories where individuals are expected to preferentially rear close relatives in order to gain inclusive fitness benefits. Here, allozyme markers are used to examine whether female Exoneura robusta individuals preferentially nest with their closest kin when given a choice of familiar previous nestmates. The results suggest these bees do not prefer kin over non-kin nestmates. Kin associations during nest founding in this species are probably due to philopatry and/or association with previously familiar nestmates.

Factors determining nest distribution in the high-mountain ant Proformica longiseta (Hymenoptera Formicidae)

In this work we study nest distribution and its determining factors in Proformica longiseta, a high-mountain ant. Four factors affecting the distribution of nests were analysed: nest density, intraspecific and interspecific competition, distribution and density of nest-site rocks, and the mode of nest founding. In general, P. longiseta shows a random distribution which is favoured by the distribution and high density of adequate nest sites, and by little or no intraspecific and interspecific competition for space. P. longiseta shows a rather stable interannual density in the study area, although there is some variation over the active period. Over-dispersion was found on only two occasions, coinciding with the highest nest density. The rocks are not a limited resource and their abundance and distribution have no effect on P. longiseta nest distribution. This species founds nests by budding, but this process appears to influence nest distribution only when plots are larger than 100 m2.

Payoffs of the two alternative nesting tactics in the African dung beetle, Scarabaeus catenatus

1. Scarabaeus catenatus is a ball‐rolling scarab in the subfamily Scarabaeinae. This species, however, makes use of two tactics for nest building: rolling and tunnelling. The tunnelling tactic differs substantially from the rolling tactic in that (1) it always involves repeated movements to and from the dung source and the nest, whereas rolling does not, and (2) it involves a shorter distance between the two sites.2. Brood‐nest founders were usually males and less often females, with about 25% adopting the rolling tactic and 75% adopting the tunnelling tactic. During nest building, the founder paired off with a scarab of the opposite sex, and they co‐operated in the work. The female made one to four brood balls from the dung in the nest, each of which contained one egg.3. Each scarab seemed to be able to employ both tactics. The tactic employed was independent of an individual's status, e.g. body size and timing of nest founding.4. The rolling tactic offered only male founders a greater nest‐defence success than the tunnelling tactic due to a lower intrusion into the rolled nest and a higher intensity of male–male fighting. The tunnelling tactic offered both male and female founders a larger number of brood balls than the rolling tactic because it enabled scarabs to take a larger amount of dung into the nest.5. The reproductive success for the two tactics was estimated from the product of nest‐defence success and the number of brood balls. As a result, the two tactics had equal fitness payoffs for males, but unequal payoffs for females.6. The results suggest that male alternation of tactics is controlled by a mixed strategy. Female alternation, however, cannot be explained by mixed strategy, alternative strategies or conditional strategy.