Intraspecific interactions in a community of arboreal nesting termites (Isoptera: Termitidae)

In coconut plantations of northern New Guinea, the arboreal nesting termite community comprises three species:Nasutitermes princeps, N. novarumhebridarum, andMicrocerotermes biroi. In orde to assess the importance of intraspecific interactions in this community, we conducted pairwise encounters between batches of individuals in the laboratory and between entire nest populations in seminantural conditions. Three levels of agonism were defined in laboratory bioassays: anagonism, moderate agonism, and strong agonism. Anagonism was observed during all control tests with homocolonical groups and in some tests with allocolonial groups of all species. Moderate agonism included initial aggressiveness that subsequently faded out, and initially passive encounters where aggression progressively built up and led to fighting. Strong agonism corresponded to initial aggressiveness and fighting. Results obtained in alboratory bioassays were consistent with bioassays in seminatural conditions. WhenNasutitermes colonies were anagonists in laboratory bioassays, their foraging trails merged without aggression in field tests.N. princeps nests that were moderately agonistic in laboratory tests fought and either continued to avoid each other or finally joined after elimination of the most aggressive individuals. The most aggressiveM. biroi andN. princeps colonies fought and their foraging trails diverged afterward. Direct attacks on alien nests were winnessed inM. biroi. In all species, anagonism occurred in 21–34% of the combinations tested, between either geographically close or distantcolonies. An exeption was a group of 112 anagonist nests ofN. princeps, which most probably constituted a supercolony. The level of agonism betweenNasutitermes colonies was constant during the wet and dry season. Termite colonies excluded each other, both intra- and interspecifically, from the coconut trees, and their territories seem distributed in a mosaic pattern. Agonism between colonies may result in the elimination of the weakest colonies or in trail divergence, maintaining this mosaic. In cotrast, lack of agonism between some colonies suggests the possibility of colony fusion and gene exchanges without nuptial flights.     

The Rules of Aggression: How Genetic,Chemical and Spatial Factors Affect Intercolony Fights in a Dominant Species,the Mediterranean Acrobat Ant Crematogaster scutellaris

Nest-mate recognition plays a key role in the biology of ants. Although individuals coming from a foreign nest are, in most cases, promptly rejected, the degree of aggressiveness towards non nest-mates may be highly variable among species and relies on genetic, chemical and environmental factors. We analyzed intraspecific relationships among neighboring colonies of the dominant Mediterranean acrobat ant Crematogaster scutellaris integrating genetic, chemical and behavioral analyses. Colony structure, parental relationships between nests, cuticular hydrocarbons profiles (CHCs) and aggressive behavior against non nest-mates were studied in 34 nests located in olive tree trunks. Bayesian clustering analysis of allelic variation at nine species-specific microsatellite DNA markers pooled nests into 14 distinct clusters, each representing a single colony, confirming a polydomous arrangement of nests in this species. A marked genetic separation among colonies was also detected, probably due to long distance dispersion of queens and males during nuptial flights. CHCs profiles varied significantly among colonies and between nests of the same colony. No relationship between CHCs profiles and genetic distances was detected. The level of aggressiveness between colonies was inversely related to chemical and spatial distance, suggesting a ‘nasty neighbor’ effect. Our findings also suggest that CHCs profiles in C. scutellaris may be linked to external environmental factors rather than genetic relationships.     

Does Distance Among Colonies and Resource Availability Explain the Intercolonial Aggressiveness in <Emphasis Type="Italic">Nasutitermes</Emphasis> aff. <Emphasis Type="Italic">coxipoensis?</Emphasis>

Aggressive behaviour can ensure animal access to local resources. To reduce constant costs in the defence of territories, species could save energy with conflicts avoiding aggression with neighbour or in situations with abundance of resources. In the present study, we analysed the effect of distance among colonies and resource availability on the aggression level and responses to chemical cues of Nasutitermes aff. coxipoensis (Holmgren) (Termitidae: Nasutitermitinae). Manipulation of resource offer was conducted in the field, where nests with different distances were kept without addition of baits (control), with addition of three or 16 sugarcane baits/nest. After 3 months, aggressiveness, linear and Y-shaped trail-following bioassays were carried out with all pairwise combinations of colonies in each treatment. Our results showed that aggressive index of N. aff. coxipoensis was affected by the resource availability. However, individuals from colonies with 0 and 3 baits/nest showed a higher number of fighting with neighbours than those from non-neighbours colonies. Termite workers from colonies without baits (control) followed shorter distance in the linear trails compared to those from colonies with addition of baits. In all treatments, there was no preference of workers in relation to the choice of chemical cues from own or other colonies. The response of intercolonial aggressiveness in N. aff. coxipoensis seems to be resource-dependent. These results may contribute to the comprehension of the use of space by N. aff. coxipoensis and could be useful to explain patterns of termite co-occurrence at different spatial scales, from local (inside the nest—e.g. cohabitation of nests by inquilines) to regional (e.g. around the nest).     

Structure and Dynamics of the Arboreal Termite Community in New Guinean Coconut Plantations21

Whereas ant mosaics have been widely recognized and described in tropical ecosystems, data on space partitioning among arboreal termite colonies are rudimentary. During a long term field study in New Guinea, the distribution of arboreal termite species in coconut plantations as well as the extent and dynamics of competition between them were investigated. The three dominant species, Mirrocerotermes biroi, Nasutitermes princeps and N. novarumhebridarum, feed on the same items but never exploit the same tree. The resulting distribution pattern is a mosaic with two peculiarities. First, some extended areas around N. princeps colonies appear unexploited, as this species practices interference competition on a wide scale, defending large territories inter- and intra-specifically. Second, interspecific relations are asymmetrical. In some plantations, large colonies of N. princeps expand their territory by destroying colonies of M. biroi, but when the pressure of N. princeps is relaxed, dense populations of colonies of M. biroi can recolonize the trees in a few years’ time. Territorial boundaries may thus change relatively fast. N. novarumhebridarum often colonizes dead trees and interferes less with the other species. These facts are consistent with each species’ reproductive investment strategy. Hypotheses are proposed to explain how the dominant species can coexist, even in long established plantations.     

Low intraspecific aggressiveness in two obligate plant-ant species

Little is known about the aggressiveness of plant-ants typically living in isolated trees nor about how that aggressiveness varies based on this isolation. Here, we examine intra- and interspecific aggressiveness between workers of two Allomerus species associated with two different myrmecophytes. In both cases, the level of intraspecific aggressiveness is very low whatever the distance separating the tested nests, while interspecific conflicts are always violent. Similar patterns of aggressiveness have been reported in various ant species, but the strictly arboreal life of Allomerus ants associated with the isolation of their adult colonies highlight different ecological conditions that might explain the lack of aggressiveness between conspecifics. Received 18 December 2007; revised 12 March 2008; accepted 13 March 2008.     

Ants under the conditions of an extremely hot summer

Specific features of ant behavior during the extremely hot summer of 2010 were studied, as well as the aftereffects of this season on simple and complex family units of ants in 2011–2012. Simultaneous studies were carried out in southern taiga (Moscow Province, Verkhnaya Klyazma myrmecological protected area) and northern taiga (Arkhangelsk Province, Pinezhskii Nature Reserve). Ants of the genus Formica responded to the extreme heat by (1) changing their foraging patterns; (2) redesigning their nests; (3) rearranging the spatial and functional structure of the colonies. They switched to a bimodal activity pattern with maxima in the morning and in the evening and a prolonged daytime intermission. Along ant roads, there were underground pavilions with dense roofs built of conifer needles and soil. Covers of the same kind appeared over root aphid colonies. The most radical improvement was the construction of a battery of brood chambers underneath the mound. The soil excavated was used for strengthening the mound and restricting the convective heat exchange between the brood chambers and the external environment. The anthill surface was covered with a smooth crusted layer of soil and fine plant debris particles, which protected the nest from the inflow of hot air from without. A portion of the inhabitants of large nests moved to newly built extensions and auxiliary nests. It was only active, non-damaged colonies that could afford these measures and thus survive the heat with minimal loss. Depressed colonies lost the major part of their brood during this time. Furthermore, small secondary colonies that emerged as the result of destructive activity of animals also failed to reassemble due to the summer heat of 2010. The months of heat were followed by a long rainy and cool period, and the colonies that had already been depressed faced critical conditions for preparing for winter. The ants were unable to accumulate lipid reserves sufficient for spring nest heating and rearing of sexuals and workers, which turned out to be an important aftereffect of the 2010 season the following year. Two first generations of workers were absent in almost all the nests. Mass oviposition in F. aquilonia commenced only late in May 2011, whereas the flight of alates did not occur at all because alates of this species are only reared in spring. Consequently, the F. aquilonia colonies had not recovered even by the end of 2012. In F. lugubris and F. polyctena, species that rear sexuals twice a year, there was only late-season alate flight, and yet it allowed their colonies to replenish the pool of ovipositing females and restore the initial colony size by the end of 2012. Other ants, represented by Lasius niger and Myrmica rubra, also modified their nests and foraging patterns during the period of heat. M. rubra suffered the most, so that its abundance and activity remained very low in 2011 as well. Only L. niger, which remained practically unscathed by the heat, successfully completed its annual cycle in 2010. Therefore, one of the 2010 year’s results was a drastic shift in the ratio of nests built by the abovementioned species in favor of L. niger.     

Community interactions between ants and arboreal-nesting termites in New Guinea coconut plantations

Summary: We investigated the incidence of inquiline ants and of arboreal-nesting ants on a community of three arboreal-nesting termites living in New Guinea coconut plantations. Inquiline ants were present in 10 % of Microcerotermes biroi nests and in 4 % of Nasutitermes princeps nests. Live termite nests inhabited by the most common inquiline ant, Camponotus sp. A, were generally left by the ant after several months. In some nests, Camponotus sp. A was observed coexisting with its host during the whole observation period (3 years). Therefore, Camponotus sp. A was apparently an opportunistic inquiline which did not affect significantly the mortality of termite colonies. The arboreal-nesting ant, Crematogaster irritabilis, was locally found occupying up to 99 % of the trees present in 1 ha plots. In such hot spots, the overall abundance of termites was approximately half that of plots devoid of Crematogaster irritabilis. The high density of Crematogaster irritabilis may be an important limiting factor for the termite assemblage, by hastening the death or hindering the establishment of arboreal termite colonies.     

Intraspecific variation in soldier defense secretions of Longipeditermes longipes (Isoptera,Nasutitermitinae)

The chemical defense secretions of major and minor soldiers of 46 colonies of the free-ranging termite Longipeditermes longipes (Isoptera, Termitidae, Nasutitermitinae) were isolated and analyzed by chromatography and spectroscopy. The colonies, collected from rainforest sites in Malaysia, showed monoterpene patterns rich in pinenes and limonene but with few qualitative differences among colonies. In marked contrast, the diterpene chemistry is highly variable that includes bicyclic (secotrinervitane), tricyclic (trinervitane), tetracyclic (rippertane), and a spirotetracyclic (longipane) skeleton. This paper presents the compositional and structural diversity, and calculates genetic distances among sympatric and allopatric colonies from a single species.     

Anti-predator behaviour in the ant Pheidole desertorum: The importance of multiple nests

The relationship between emigrations and anti-predator behaviour in the ant Pheidole desertorum was investigated in the field. Frequent emigrations in P. desertorum result in the colony having multiple nests, of which only one is occupied at a time. A field procedure demonstrated that the ants quickly find and enter the unoccupied nests after a nest evacuation caused by the army ant Neivamyrmex nigrescens. When colonies were denied access to their unoccupied nests after an evacuation, survival of the brood and alates was significantly lower than in colonies allowed access to all their nests. These results suggest that emigrations in P. desertorum are part of a defence strategy against army ants.     

Intraspecific aggression and the colony structure of the invasive ant Myrmica rubra

1. Patterns of aggression between ants from different nests influence colony and population structure. Several species of invasive ants lack colony boundaries over large expanses, forming ‘supercolonies’ with many nests among which workers can move without encountering aggression. 2. Bioassays of aggression were used to determine the colony structure of the invasive ant Myrmica rubra (L.) at eight sites in Massachusetts, the state where the species was first discovered in North America. To improve the ability to distinguish systematic patterns from background variability in aggressiveness, a repeated‐measures design was used and replicate assays for each pair of nests were conducted. 3. Aggressive responses showed that populations at all sites consisted of multiple distinct colonies. Patterns of aggression were repeatable and transitive, with few exceptions. Colonies were identified as clusters of nests whose workers showed little to no aggression towards one another but were aggressive towards conspecifics from more distant nests. 4. The degree of aggression varied considerably among different colony pairs but did not depend in any consistent way on the distance of separation or on whether colonies were neighbours. 5. Territories of neighbouring colonies abutted, indicating that they were restricted by intraspecific competition. Mapped territories ranged in size from 0.03 to 1.2 ha, but colonies at the study sites have not undergone the enormous expansions seen in introduced populations of some other species of invasive ants, and neighbouring colonies compete locally.     

The Role of Non-Foraging Nests in Polydomous Wood Ant Colonies

A colony of red wood ants can inhabit more than one spatially separated nest, in a strategy called polydomy. Some nests within these polydomous colonies have no foraging trails to aphid colonies in the canopy. In this study we identify and investigate the possible roles of non-foraging nests in polydomous colonies of the wood ant Formica lugubris. To investigate the role of non-foraging nests we: (i) monitored colonies for three years; (ii) observed the resources being transported between non-foraging nests and the rest of the colony; (iii) measured the amount of extra-nest activity around non-foraging and foraging nests. We used these datasets to investigate the extent to which non-foraging nests within polydomous colonies are acting as: part of the colony expansion process; hunting and scavenging specialists; brood-development specialists; seasonal foragers; or a selfish strategy exploiting the foraging effort of the rest of the colony. We found that, rather than having a specialised role, non-foraging nests are part of the process of colony expansion. Polydomous colonies expand by founding new nests in the area surrounding the existing nests. Nests founded near food begin foraging and become part of the colony; other nests are not founded near food sources and do not initially forage. Some of these non-foraging nests eventually begin foraging; others do not and are abandoned. This is a method of colony growth not available to colonies inhabiting a single nest, and may be an important advantage of the polydomous nesting strategy, allowing the colony to expand into profitable areas.     

A mimetic nesting association between a timid social wasp and an aggressive arboreal ant

In French Guiana, the arboreal nests of the swarm-founding social wasp Protopolybia emortualis (Polistinae) are generally found near those of the arboreal dolichoderine ant Dolichoderus bidens. These wasp nests are typically protected by an envelope, which in turn is covered by an additional carton ‘shelter’ with structure resembling the D. bidens nests. A few wasps constantly guard their nest to keep D. bidens workers from approaching. When alarmed by a strong disturbance, the ants invade the host tree foliage whereas the wasps retreat into their nest. Notably, there is no chemical convergence in the cuticular profiles of the wasps and ants sharing a tree. The aggressiveness of D. bidens likely protects the wasps from army ant raids, but the ants do not benefit from the presence of the wasps; therefore, this relationship corresponds to a kind of commensalism.     

Nesting Biology of the Bee Hoplitis princeps (Morawitz) (Hymenoptera,Megachilidae) in Crimea

Bionomics of Hoplitis princeps (Morawitz) was studied in Crimea. The species is confined to coastal psammophytic habitats and has one generation per year. Females nest in sand and excavate burrows near roots of various plants. The nests consist of 1 or rarely 2 cells arranged either side by side or one by one; one nest with 3 cells made by two different females was also discovered. The building material was masticated leaves of Astragalus varius subsp. eupatoricus Sytin; the provision was pollen and nectar from the same plant species. The cleptoparasitic bee Stelis aculeata Morawitz was discovered in the nests. The differences in the nest structure between H. princeps and a closely related species H. fulva (Eversmann) are discussed. The need for conservation of the H. princeps habitat in Crimea is substantiated.

     

Nest-specific composition of the trail pheromone of the stingless bee Trigona corvina within populations

Social insects have evolved highly developed communication systems, enabling them to coordinate complex interactions in their colonies. Pheromones play a major role in the coordination of many tasks. In Trigona corvina, a stingless bee that occurs in Central America, foragers use pheromones produced in their labial glands to scent mark solid substrates between a food source and their nest. Newly recruited bees subsequently follow these scent marks until they reach the food source. A recent study has revealed nest-specific differences in the composition of these trail pheromones in colonies of T.?corvina, suggesting that pheromone specificity may serve to avoid competition between foragers from different nests. However, the nests used in this study came from different populations and their foragers certainly never met in the field (Jarau et al., 2010). The aim of the present study was to investigate whether differences in the trail pheromones of foragers from different nests can also be found between neighbouring colonies within populations. We analysed the composition of trail pheromones from labial gland secretions extracted from workers from nine colonies collected at three different populations in Costa Rica. The differences in pheromone composition were even more distinct between neighbouring nests within a population than between nests of different populations. This finding corroborates the hypothesis that nest specificity of trail pheromones serves to communicate the location of a food source exclusively to nestmates, thereby avoiding intraspecific competition at resources. Resource partitioning by avoiding conspecific non-nestmates is particularly adaptive for aggressive bee species, such as T. corvina.     

Relationship between the age of Vespa velutina workers and their defensive behaviour established from colonies maintained in the laboratory

As the structural bases of insect societies are essential to colony survival, nests must be protected from predation. Nest defence behaviours are among the most important roles assigned to worker members. However, in hornet societies, temporal polyethism (age-dependent division of labour among workers) is assumed to be weak. Few studies have investigated this phenomenon, probably because hornet nests are aggressively defended and dangerous to approach. In the present study, we propose a method for rearing nests of Vespa velutina, a species newly introduced into Europe. This method enables the handling of hornets, and thus the design of experiments. By marking all newly emerged hornets, we recorded aggressiveness in workers of different ages from three captive colonies. We observed that nest defence behaviour in V. velutina depends on the age of the workers. Nest defence appears to be mediated by the queen, probably through pheromones that promote nest organization. We also identified a previously unreported but important behaviour in V. velutina that workers are aggressive towards male hornets. This behaviour might be a strategy to avoid inbreeding. Collectively, our results provide new research perspectives for the management of social hymenopteran predators.     

Queen-worker conflict and social parasitism in bumble bees (Hymenoptera: Apidae)

Psithyrus ashtoni (Hymenoptera: Apidae) is an obligate, workerless bumble, bee social parasite which invades nests of Bombus affinis. Although parasites are limited by host worker defence to invading very small colonies, there is considerable flexibility in the way parasites control host brood bionomics once they are accepted inside the nest. A study of 46 parasitized and 22 non-parasitized laboratory colonies of B. affinis showed that P. ashtoni females cohabited with host queens and workers while the worker force increased, but not to the maximum normally achieved in non-parasitized nests. While colony reproductive success was correlated with the number of workers reared, parasites risked being killed or ejected from the comb by workers, after the queen had lost dominance. Host bees usually succeeded in rearing offspring, and Psithyrus reproductive success was related to the ability of parasites to control proportional investment in the two species. In addition to displacing P. ashtoni females, host bees ate the eggs of parasites and ejected their larvae. These behaviours were also exhibited by workers in the later stages of development of non-parasitized colonies. These results indicate that social parasites are at least partially subject to the conflict of genetic self-interest between the queen and her workers which is believed to influence the control of reproductive investment in haplodiploid Bombus societies.     

Multiple queens in founding colonies of the neotropical ant Pachycondyla striata Smith (Formicidae: Ponerinae)

In social insects, the typical mode of colony foundation occurs when a single queen is inseminated by a male and establishes a new colony, although we can find interspecific and intraspecific variations in queen number and queen-mating frequencies in a single colony. This study aimed to verify the queen number in Pachycondyla striata (Smith) colonies and to evaluate the level of aggressiveness among workers. We collected 14 colonies of P. striata. The behaviors of individuals from five multiple-queen colonies maintained in laboratory were studied by the method of scan sampling. In order to evaluate aggressiveness, dyadic encounters among heterocolonial and homocolonial workers were performed. The results showed that colonies of P. striata can have two or more mated queens (polygynous colonies) besides to monogynous ones (colony containing one queen). Because in polygynous colonies the number of workers was relatively low, such colonies could represent colonies in the foundation phase that characterize a pleometrosis state. In fact, ovarian development analysis from queens showed that the number of queens in the colonies seemed to be unstable. Despite a few cases of oophagy (egg cannibalism), social hierarchy among queens is unclear in comparison to other Pachycondyla species. In addition, aggressiveness increased with distance among nests. Nearby colonies (less than 1 m apart) showed a low level of aggressiveness, suggesting the presence of polydomy, that is, a unique colony can occupy multiple nests. Polygyny associated to polydomy in founding colonies may confer benefits on growth and dispersion of colonies in the studied environments.     

Brood reduction in temperate and sub-tropical ospreys

Summary In an effort to understand patterns and causes of nestling loss in Ospreys (Pandion haliaetus), I studied brood reduction in three eastern U.S. Osprey colonies during 1978 and 1979. The colonies, located in Florida Bay (1) and on coastal Long Island, N.Y. (2), differed in the average daily amount of food delivered to nestlings; Florida nests received 43% and 11% less fish per day than nests in the two N.Y. colonies, largely because latitude and season restricted day length and thus foraging time for the winter-breeding Florida Ospreys. Increased distance from stable food sources accounted for the lower rate of feeding at one of the N.Y. colonies. Variation in clutch size in the three colonies reflected differences in latitude more than in food availability; average clutch sizes in Long Island were larger than Florida clutches by 0.5 of an egg, but were similar to each other and to those in other northeastern U.S. Osprey populations.Increased nestling loss coincided with reduced food delivery rates and, in food stressed colonies, this loss was 2–3 times greater than any recorded for Ospreys. Starvation was the primary cause of nestling death, with mortality concentrated on third chicks, which hatched on average 3.9 d later and from eggs 5.6% smaller than chicks hatching first. Sibling aggression accounted for the preferential feeding of older nestmates,but only in colonies or nests where food was limited. Aggressive chicks nearly always stopped fighting after being fed. This behavior provided a reversible mechanism for controling brood reduction that was based on nutrition. Growth rates of young measured during the first half of the growth period were more variable between colonies than within nests. This is interpreted as reflecting both the differences in colony food delivery rates as well as the evolutionary pressures of sibling competition to equalize the growth of nestmates.     

Nest relocation in the slavemaking ants Formica subintegra and Formica pergandei: a response to host nest availability that increases raiding success

Social insects typically occupy spatially fixed nests which may thus constrain their mobility. Nevertheless, colony movements are a frequent component of the life cycle of many social insects, particularly ants. Nest relocation in ants may be driven by a variety of factors, including nest deterioration, seasonality, disturbances, changes in microclimate, and local depletion of resources. The colony movements of slavemaking ants have been noted anecdotally, and in recent studies such relocations were primarily attributed to nest deterioration or shifts to overwintering locations. In this study we explore nest relocations in large colonies of formicine slavemakers which occupy stable and persistent earthen nest mounds. We investigate the hypothesis that colony relocations of these slavemakers are best explained by efforts to improve raiding success by seeking areas of higher host availability. Five summers of monitoring the raiding behavior of 11–14 colonies of the slavemakers Formica subintegra and Formica pergandei revealed relatively frequent nest relocations: of 14 colonies that have been tracked for at least three of 5 years, all but one moved at least once by invading existing host nests. Movements tended to occur in the middle of the raiding season and were typically followed by continued raiding of nearby host colonies. Spatial patterns of movements suggest that their purpose is to gain access to more host colonies to raid: the distance moved is typically farther than the mean raiding distance before the move, which may indicate an effort to escape their local neighborhood. Furthermore, the mean distance of raids after relocation is shorter than the distance before relocation. For many slavemaking ant colonies, particularly those on the verge of relocating, raiding distance increased as the raiding season progressed. In addition, movements tended to be toward areas of higher local host density. Nest relocation is likely an important component of the ecology of slavemaking ants that contributes to the dynamic nature of their interaction with the host ant population.     

Seasonal patterns of egg production in field colonies of the termite <Emphasis Type="Italic">Reticulitermes speratus</Emphasis> (Isoptera: Rhinotermitidae)

This is the first report on the annual egg production patterns in mature termite colonies in the field. Data on the seasonal patterns of egg production in field colonies are very important for understanding the annual colony growth schedule, resource allocation, and population dynamics of the termites. However, collecting the eggs from a sufficient number of colonies is extremely difficult in Reticulitermes termites because their multiple-site nesting makes it difficult to find the reproductive center of the colonies. Here, we first show the seasonal pattern of egg production in the subterranean termite Reticulitermes speratus by collecting the reproductive center of ten colonies each month from April through October. We had to destructively examine dozens of nests to find eggs from enough field colonies each month. Mature field colonies began to produce eggs in late May, soon after the swarming season, and the egg production rate (EPR) reached its maximum in early July. The eggs hatched until late October. The EPR was significantly correlated with the average monthly temperature. Additional investigation of the egg distributions in the nests showed that most eggs were kept around the royal cell, which contained the reproductives. The largest colony had 109 supplemental queens and 94,023 eggs, suggesting that each queen produced an average of 24.7 eggs per day, based on the known mean hatching period of an inseminated egg of 34.95±0.12 (SE) days.