Sexual Brood Research Articles

Sex-specific growth of nestlings of the Whiskered Tern Chlidonias hybrida, a species with sexual size dimorphism and female brood desertion

The growth of a nestling during the first few weeks after hatching is crucial for its further life and is sensitive to the conditions experienced during this period. Among species exhibiting Sexual Size Dimorphism (SSD), one of the most important factors influencing growth parameters is the sex of the hatchlings. In this study, we tested whether sex and other factors (hatching date, egg volume, brood size and sex ratio in the brood) were related to sex on growth parameters in Whiskered Tern (Chlidonias hybrida) chicks. In this species, males are 3%–10% larger than females when adult. At hatching, only the total head length was greater in males than females, but values close to fledging were greater for most of the measured parameters (3%—tarsus, 4%—total head length, 13%—body mass). Moreover, the maximum body mass growth rate was higher in males but females entered the rapid growth phase somewhat earlier and achieved the final total head length sooner. Nestling growth was also significantly correlated with brood size and hatching date, though mostly negatively (e.g. lower tarsus and total head length asymptote, lower maximum growth rate and growth rate constant of total head length in bigger and later clutches). The occurrence of SSD during the nestling period and higher maximum growth rate of body mass in males indicates that the costs of raising sons may be higher than of raising daughters. However, we did not find evidence of either sex-biased nestling mortality or skewed sex ratio.

Caste-dependent brood retrieval by workers in the ant Formica exsecta

The ability to distinguish friends from foe is a widespread phenomenon among social animals. In ants, recognition of intruders is important for the maintenance of colony integrity and survival. Intruders are typically adult, but the acceptance of non-nestmate brood could result in severe fitness costs, depending on the caste of the brood. Accepting non-nestmate worker brood may not carry a cost, as they should not drain resources of the adoptive colony but may instead add to the workforce. Sexual brood, however, would typically not contribute to colony performance, yet require resources, and should thus be rejected. Here, we tested whether workers of the narrow-headed ant, Formica exsecta, which strongly discriminate between adult nestmates and non-nestmates, also discriminate between nestmate and non-nestmate pupae. Furthermore, we investigated whether the caste of the brood (workers/sexuals) affects discrimination. We carried out analysis of surface chemicals to investigate whether the chemical distance between colonies was associated with the propensity to accept non-nestmate pupae. We show that worker pupae were retrieved irrespective of their origin, whereas nestmate sexual pupae were retrieved at a slightly higher rate than non-nestmates. Our chemical data, however, suggest that both the reproductive and the worker brood carry sufficient chemical information for discrimination, as they both express colony signatures. However, this information is acted upon only in the case of sexual brood. Our results thus suggest that workers selectively capitalize on the chemical information in agreement with fitness predictions, albeit to a lower extent than during discrimination between adult individuals.

Caste regulation in the ant Monomorium pharaonis (L.) with emphasis on the role of queens

The role of queens in caste regulation of pharaoh ant colonies has been puzzling biologists for decades. Worker-fertility-inhibiting pheromones are lacking, as workers of this polygynous species are completely sterile. Yet, fertile queens possess inhibiting powers over the production of new gynes and males. Here we test whether queen-produced cuticular compounds or factors related to eggs affect sexual brood survival. Additionally, the influence of dead queens and virgin queens is investigated. Hereto, subcolonies consisting of workers, mixed worker brood and sexual larvae are treated with eggs, living queens, dead queens, virgin queens or queen extracts. The extracts are prepared with (phosphate-buffered saline), hexane or acetone and applied on either glass dummies (chemical cue only) or queen dummies (visual and chemical cue). Subsequently, the number of surviving larvae and their sex ratio are recorded. Only treatments containing living queens show a significant drop in eclosing sexual larvae. In all other treatments the survival of sexual larvae is unaffected. Several hypotheses are discussed involving competition for ‘royal’ food between queens and sexual larvae, food deprival of sexual larvae by workers or cannibalism of sexual brood by queens and/or workers. Although brood cannibalism seems the most valid explanation, as it allows recycling the energy used for brood production throughout the colony, our data could not sufficiently support this hypothesis. Therefore, additional research on different aspects of caste regulation is required to fully understand its mechanism in pharaoh ants.

Acromyrmex charruanus: a new inquiline social parasite species of leaf-cutting ants

Social parasites exploit the colony resources of social species to secure their own survival and reproduction. Social parasites are frequently studied as models for conflict and cooperation as well as for speciation. The eusocial Hymenoptera harbor a diverse array of socially parasitic species with idiosyncratic life history strategies, but it is probably in the ants where social parasites are most speciose and have evolved the highest degrees of morphological and behavioral specialization. In the fungus-growing ants, a total of five obligate social parasites are known: four species are parasites of leaf-cutting ants and one species parasitizes a primitive fungus-growing ant species in the genus Mycocepurus. Here we describe a new species of socially parasitic leaf-cutting ant, Acromyrmex charruanus sp. nov., from Uruguay, and we report initial observations on the parasite’s life history as well as on the morphological and behavioral adaptations related to the inquiline syndrome. Our observations suggest that Acromyrmex charruanus is an obligate inquiline social parasite of the thatch-mound-building, leaf-cutting ant Acromyrmex heyeri. Acromyrmex charruanus appears to be tolerant of the host, producing sexual offspring in the presence of the A. heyeri host queen. Queens of A. charruanus appear to reproduce semelparously and sexual offspring are produced during the austral fall (February), which differs significantly from the mating biology of the host species, which reproduces during the southern-hemisphere spring (October–December). We suggest that the diametrically opposed mating seasons of parasite and host might be adaptive, allowing the parasite to avoid competition for resources with the host sexual brood. The morphological and behavioral adaptations of A. charruanus accord with characters observed to arise early during the evolution of other ant inquiline parasite species, and so far we have no evidence for the existence of a worker caste in A. charruanus. Further field studies and behavioral experiments need to confirm our first observations and explore A. charruanus’s behavioral ecology, evolution, and life history in more detail.

Thermoregulation strategies in ants in comparison to other social insects, with a focus on red wood ants ( Formica rufagroup).

Temperature influences every aspect of ant biology, especially metabolic rate, growth and development. Maintenance of high inner nest temperature increases the rate of sexual brood development and thereby increases the colony fitness. Insect societies can achieve better thermoregulation than solitary insects due to the former's ability to build large and elaborated nests and display complex behaviour. In ants and termites the upper part of the nest, the mound, often works as a solar collector and can also have an efficient ventilation system. Two thermoregulatory strategies could be applied. Firstly the ants use an increased thermal gradient available in the mound for brood relocation. Nurse workers move the brood according to the thermal gradients to ensure the ideal conditions for development. A precise perception of temperature and evolution of temperature preferences are needed to make the correct choices. A second thermoregulatory strategy used by mound nesting ants is keeping a high temperature inside large nests. The unique thermal and insulation properties of the nest material help to maintain stable conditions, which is the case of the Wood ant genus Formica. Ants can regulate thermal loss by moving nest aggregation and alternating nest ventilation. Metabolic heat produced by ant workers or associated micro organisms is an important additional source of heat which helps to maintain thermal homeostasis in the nest.

Seasonal shifts in macronutrient preferences in supercolonies of the invasiveYellowCrazyAntAnoplolepis gracilipes(Smith, 1857) (Hymenoptera:Formicidae) onChristmasIsland,IndianOcean

AbstractThe timing and duration of sexual brood production in ants can affect their rate of spread and colony growth. Because protein is key to larval growth, queen survival and fecundity, macronutrient collection by foraging workers is expected to favour protein prior to and throughout gyne production. However, food preference driven by the production of gynes may be overridden by a preference associated with worker production, especially if investment in workers vastly outweighs that of sexual brood and workers are produced on different schedules. Food preferences alone may not indicate the availability of that food type in the environment. OnChristmasIsland,IndianOcean, sexual brood of the invasiveYellowCrazyAnt,Anoplolepis gracilipes(Smith, 1857), was produced annually during a single period associated with the onset of the wet season. However, workers were produced continuously throughout the year and colony investment to worker production measured by the standing biomass of eggs, larvae and pupae, typically exceeded 98%. High, aseasonal investment in worker production, together with aseasonal worker activity, would suggest that there should be no seasonal preference shown by workers at food stations containing both protein and carbohydrate. However, workers showed a preference for one food type over another on 46 of 61 occasions at one site and on 31 of 41 occasions at another. When a preference was shown, it was predominantly for protein‐rich food during the dry season and almost always for carbohydrate at the onset and during the wet season. We suggest that these preferences reflect seasonal shortages in key resources when invertebrates (protein‐rich) are scarce during the dry season and honeydew from scale insects (carbohydrate) is depleted by rain during the wet season. OnChristmasIsland, timing and duration of the dry season preference for protein has been exploited by the control program forA. gracilipessupercolonies, which deploys a toxin in a proteinaceous bait matrix during dry periods.

Thermoregulation strategies in ants in comparison to other social insects, with a focus on Formica rufa

Temperature influences every aspect of ant biology, especially metabolic rate, growth and development. Maintenance of high inner nest temperature increases the rate of sexual brood development and thereby increases the colony fitness. Insect societies can achieve better thermoregulation than solitary insects due to the former’s ability to build large and elaborated nests and display complex behaviour. In ants and termites the upper part of the nest, the mound, often works as a solar collector and can also have an efficient ventilation system. Two thermoregulatory strategies could be applied. Firstly the ants use an increased thermal gradient available in the mound for brood relocation. Nurse workers move the brood according to the thermal gradients to ensure the ideal conditions for development. A precise perception of temperature and evolution of temperature preferences are needed to make the correct choices. A second thermoregulatory strategy used by mound nesting ants is keeping a high temperature inside large nests. The unique thermal and insulation properties of the nest material help to maintain stable conditions, which is the case of the Wood ant genus Formica. Ants can regulate thermal loss by moving nest aggregation and alternating nest ventilation. Metabolic heat produced by ant workers or associated micro organisms is an important additional source of heat which helps to maintain thermal homeostasis in the nest.

The value of oviposition timing, queen presence and kinship in a social insect

Reproductive cooperation confers benefits, but simultaneously creates conflicts among cooperators. Queens in multi-queen colonies of ants share a nest and its resources, but reproductive competition among queens often results in unequal reproduction. Two mutually non-exclusive factors may produce such inequality in reproduction: worker intervention or queen traits. Workers may intervene by favouring some queens over others, owing to either kinship or queen signals. Queens may differ in their intrinsic fecundity at the onset of oviposition or in their timing of the onset of oviposition, leading to their unequal representation in the brood. Here, we test the role of queen kin value (relatedness) to workers, timing of the onset of oviposition and signals of presence by queens in determining the maternity of offspring. We show that queens of the ant Formica fusca gained a significantly higher proportion of sexuals in the brood when ovipositing early, and that the presence of a caged queen resulted in a significant increase in both her share of sexual brood and her overall reproductive share. Moreover, the lower the kin value of the queen, the more the workers invested in their own reproduction by producing males. Our results show that both kinship and breeding phenology influence the outcome of reproductive conflicts, and the balance of direct and indirect fitness benefits in the multi-queen colonies of F. fusca.

Reproductive competition in the bumble-bee Bombus terrestris: do workers advertise sterility?

Reproductive competition in social insects is generally mediated through specific fertility pheromones. By analysing Dufour's gland secretion in queens and workers of Bombus terrestris under varying social conditions, we demonstrate here that the volatile constituents of the secretion exhibit a context-dependent composition. The secretion of egg-laying queens is composed of a series of aliphatic hydrocarbons (alkanes and alkenes), while that of sterile workers contains in addition octyl esters, dominated by octyl hexadecanoate and octyl oleate. These esters disappear in workers with developed ovaries, whether queenright (QR) or queenless (QL), rendering their secretion queen-like. This constitutes an unusual case in which the sterile caste, rather than the fertile one, possesses extra components. Individually isolated (socially deprived) workers developed ovaries successfully, but failed to oviposit, and still possessed the octyl esters. Thus, whereas social interactions are not needed in order to develop ovaries, they appear essential for oviposition and compositional changes in Dufour's gland secretion (ester disappearance). The apparent link between high ester levels and an inability to lay eggs lends credence to the hypothesis that these esters signal functional sterility. We hypothesize that by producing a sterility-specific secretion, workers signal that 'I am out of the competition', and therefore are not attacked, either by the queen or by the reproductive workers. This enables proper colony function and brood care, in particular sexual brood, even under the chaotic conditions of the competition phase.

Complex sociogenetic organization and the origin of unrelated workers in a eusocial sweat bee, Lasioglossum malachurum

Sweat bees (Halictidae) exhibit great interspecific and intraspecific diversity in their social organisation, yet there is remarkably little information on the sociogenetic organisation of any species. Lasioglossum malachurum is a eusocial sweat bee with an annual lifecycle that exhibits considerable variation in its social organisation across its wide geographic range from northern to southern Europe. We collected all adults from 31 L. malachurum nests at Eichkogl, Austria, near the latitudinal centre of its distribution, and genotyped 148 workers using 5 highly variable microsatellite loci developed for this species. Nests were often queenless (48% of nests) during the second phase of worker activity, when colonies were provisioning the sexual brood. Pedigree reconstruction and estimates of nestmate genetic relatedness demonstrated that nests often (32% of nests) contained alien workers, probably as a result of worker drifting from their natal to a foreign nest. Queen effective mating frequency was variable (harmonic mean me = 1.24), but sometimes high (maximum 2.7). These data demonstrate that nests of L. malachurum do not have a classical eusocial sociogenetic organisation (monogyny, monandry) and thereby pose a challenge to exclusively relatedness based arguments for the evolution of eusociality in the taxon.

Thermoregulatory brood transport in the fire ant, Solenopsis invicta

Nest structure in ants is often designed to optimize the colony’s ability to thermoregulate, and this specialization is most highly developed in mound-building ant species. Solenopsis invicta invest a large amount of energy in building mounds and transporting their brood up and down in their nests as a means of thermoregulation. Because few ant species build true mounds, we wanted to determine the effectiveness of these mounds in harvesting solar heat as well as to distinguish what factors (temperature vs. circadian rhythm) govern where fire ants place their brood in the mound and when they place it. We measured temperature patterns in the mound over several days at different depths and under different conditions (under direct sunlight or shade), and then conducted a series of field experiments to manipulate the orientation and time of heating. On cool mornings in spring or fall, surface temperatures of the mound rise at the fastest rate on the side receiving the most direct sunlight (usually the south side). This heating causes a temperature gradient through different depths in the mound, and shows little difference from outside ground temperature at a depth greater than ~40 cm inside the nest. In the morning, fire ants move their brood up into the mound on the side most directly heated, and when temperatures exceed optimal (~32°C) they move their brood down the temperature gradient to lower depths in the nest. In addition to this, mound temperature does not only increase due to direct sunlight, but temperature also increases higher than ground temperatures when the mound is in the shade due to its low specific heat. Experiments in which sunlight was mirrored to the normally shaded side of the mound, or when mounds were heated at night, revealed that S. invicta primarily track temperature patterns and do not rely on behavioral habits or circadian rhythms for the thermoregulatory transport of their brood. When mounds were shaded, S. invicta brood was evenly distributed directly under the surface of the mound rather than aggregating towards a specific side. The fire ant mound is important for thermoregulation because, compared to moundless subterranean nests, it absorbs heat more rapidly both in direct sunlight and shady conditions. Temperature tracking within the nest is key to understanding thermoregulatory placement of fire ant brood, as well as insight into the production of sexual brood and reproduction.

Taxonomy, distribution and nesting biology of species of the genus Dolichovespula (Hymenoptera, Vespidae)

AbstractThe genus Dolichovespula consists provisionally of 18 species. Aerial nest site characteristics have been established for seven species but most of these species show flexibility in their choice of nest sites and two species often nest in shallow subterranean situations. Colony length is usually just over 3 months to approximately 4 months, but at lower latitudes may extend to more than 5 months. Mature colony size is usually approximately 1000 cells with more large cells than small cells. Parasitized colonies have a shorter colony cycle and smaller colonies. During the production of the sexual brood the larva/worker ratio reaches its lowest value of approximately 2.5 larvae per worker. Colonies often have upper mixed combs of small and large cells. Total adult production is usually less than 2000 adults. Colonies of D. arenaria and D. norwegica may specialize in mainly producing males or queens. Variations in mature colony size and production of queens is probably a consequence of the number of workers present, particularly early in the colony cycle.

The Natural History of the Arboreal Ant, Crematogaster ashmeadi

The arboreal ant, Crematogaster ashmeadi Emery (Hymenoptera: Formicidae), is the most dominant arboreal ant in the pine forests of the coastal plain of northern Florida. The majority of pine trees harbor a colony of these ants. The colonies inhabit multiple chambers abandoned by bark-mining caterpillars, especially those of the family Cossidae, in the outer bark of living pines. They also inhabit ground level termite galleries in the bark, often locating the queen in galleries. The density of chambers and ants is highest in the base of the tree and drops sharply with height on the trunk. Because chambers are formed in the inner layer of bark, they gradually move outward as more bark layers are laid down, eventually sloughing off the tree's outer surface. Chambers have a mean lifetime of about 25 yr. The abundant chambers in pine bark are excavated by a small population of caterpillars and accumulate over decades. Ant colonies also inhabit abandoned galleries of woodboring beetles in dead branches in the crowns of pines. Because newly mated queens found colonies in abandoned woodboring beetle galleries in the first dead branches that form on pine saplings, C. ashmeadi is dependent on cavities made by other insects throughout its life cycle, and does little if any excavation of its own. Mature colonies nest preferentially in chambers greater than 10 cm2 in area, a relatively rare chamber size. In natural pine forests, this does not seem to limit the ant's populations.Founding queens containabout 50% fat and lose about half of their dry weight during the claustral period, converting approximately half of this lost weight into progeny. The claustral period is about 40 to 50 days at 27°C. Mature colonies contain several tens of thousands of workers (est. up to 80,000), and have a life expectancy of 10 to 15 years. Each colony occupies an entire tree, and sometimes two trees if they are close together. Within a colony, there is a single queen capable of laying up to 450 eggs/day during the warm season. Such queens weigh 12 to 18 mg, have 50 to 60 active ovarioles and 120 to 600 vitellogenic oocytes in their ovaries.Mature colonies begin producing sexual brood in late April or early May. Sexual adults are present from late May through June. Mating flights commence in June and most sexuals have left their natal nests by late July. Female sexuals are an especially large investment; the energetic content of a single, flight-ready female sexual is almost 20 times that of a worker. The newly mated queen sequesters a mean of 2.64 × 106 sperm in her spermatheca, a supply that should last her for 16 years at the observed reproductive rates.

Spatial patterns, temporal variability, and the role of multi‐nest colonies in a monogynous Spanish desert ant

Abstract 1. The colonies of the Spanish desert ant Cataglyphis iberica are polydomous. This study describes the temporal and spatial patterns of the polydomy in this species at two different sites, and presents analyses of its role in reducing the attacks of the queen over sexual brood, and in allowing better habitat exploitation.2. The spatial distribution of nests was clumped while colonies were distributed randomly. Mean nearest neighbour distance ranged from 3.4 to 7.0 m for nests and from 12.3 to 14.1 m for colonies. Distance of foragers searching for food varied among nests: mean values were between 6.1 and 12.6 m.3. At both sites, the maximum number of nests per colony occurred in summer, during the maximum activity period of the species. Colonies regrouped at the end of this period but overwintered in several nests.4. Nest renewal in C. iberica colonies was high and showed great temporal variability: nests changed (open, close, re‐open) continuously through the activity season and/or among years. The lifetime of up to 55% of nests was only 1–3 months.5. Polydomy in C. iberica might decrease the interactions between the queen and the sexual brood. In all colonies excavated just before the mating period, the nest containing the queen did not contain any virgin female. Females were in the queenless nests of the colony.6. The results also suggest that polydomous C. iberica colonies may enhance habitat exploitation because foraging activity per colony increases with nest number. The relationship between total prey input and foraging efficiency and number of nests per colony attains a plateau or even decreases after a certain colony size (four to six nests). This value agrees with the observed mean number of nests per colony in C. iberica.

The natural history of the arboreal ant, Crematogaster ashmeadi.

The arboreal ant, Crematogaster ashmeadi Emery (Hymenoptera: Formicidae), is the most dominant arboreal ant in the pine forests of the coastal plain of northern Florida. The majority of pine trees harbor a colony of these ants. The colonies inhabit multiple chambers abandoned by bark-mining caterpillars, especially those of the family Cossidae, in the outer bark of living pines. They also inhabit ground level termite galleries in the bark, often locating the queen in galleries. The density of chambers and ants is highest in the base of the tree and drops sharply with height on the trunk. Because chambers are formed in the inner layer of bark, they gradually move outward as more bark layers are laid down, eventually sloughing off the tree's outer surface. Chambers have a mean lifetime of about 25 yr. The abundant chambers in pine bark are excavated by a small population of caterpillars and accumulate over decades. Ant colonies also inhabit abandoned galleries of woodboring beetles in dead branches in the crowns of pines. Because newly mated queens found colonies in abandoned woodboring beetle galleries in the first dead branches that form on pine saplings, C. ashmeadi is dependent on cavities made by other insects throughout its life cycle, and does little if any excavation of its own. Mature colonies nest preferentially in chambers greater than 10 cm(2) in area, a relatively rare chamber size. In natural pine forests, this does not seem to limit the ant's populations. Founding queens contain about 50% fat and lose about half of their dry weight during the claustral period, converting approximately half of this lost weight into progeny. The claustral period is about 40 to 50 days at 27 degrees C. Mature colonies contain several tens of thousands of workers (est. up to 80,000), and have a life expectancy of 10 to 15 years. Each colony occupies an entire tree, and sometimes two trees if they are close together. Within a colony, there is a single queen capable of laying up to 450 eggs/day during the warm season. Such queens weigh 12 to 18 mg, have 50 to 60 active ovarioles and 120 to 600 vitellogenic oocytes in their ovaries. Mature colonies begin producing sexual brood in late April or early May. Sexual adults are present from late May through June. Mating flights commence in June and most sexuals have left their natal nests by late July. Female sexuals are an especially large investment; the energetic content of a single, flight-ready female sexual is almost 20 times that of a worker. The newly mated queen sequesters a mean of 2.64 x 10(6) sperm in her spermatheca, a supply that should last her for 16 years at the observed reproductive rates.

Role of resource availability on sex, caste and reproductive allocation ratios in the Argentine antLinepithema humile

SummaryResource availability has long been recognized to influence reproductive decisions in eusocial Hymenoptera. We analysed how protein availability affects the number and the weight of worker, male and queen pupae, as well as its effect on sex, caste and reproductive allocation ratios, in the Argentine antLinepithema humileMayr. Colonies were maintained on diets with three levels of access to proteins: no protein, intermediate and high levels of proteins.There was no significant difference between the intermediate and high levels of protein in the number of queen, male and worker pupae produced. Similarly, the intermediate vs. high levels of protein treatments did not differ with regard to the weight of pupae nor the relative investments between the three castes. This suggests that, inL. humile, there could be a threshold over which additional amounts of protein have no more effect on reproductive allocation or the size of the individuals produced.The main effect of protein supplementation was to increase the number of sexual pupae (queens and males) produced. By contrast, the number of workers produced remained unaffected by the level of proteins. The higher investment in sexuals in nests supplemented with proteins resulted in a significantly higher proportion of queen pupae among females, as well as higher proportion of males among all individuals produced in these nests. However, the proportion of males among sexuals (numerical sex ratio) was not significantly different between supplemented and unsupplemented nests.Workers reacted to a higher protein diet not only by rearing more sexuals, but also by producing larger individuals of the three castes.The proportional investment allocated to queens among females was higher in the protein‐supplemented treatments. Protein‐supplemented nests also invested proportionally more into sexual production than unsupplemented nests. However, whereas increased sexual production is generally associated with a more female‐biased sex investment ratio in ants, sex allocation ratio was not significantly different between supplemented and unsupplemented nests.Overall, our results support the view that protein availability influences the proportion of brood that is culled, with workers eliminating a significant proportion of both the male and female sexual brood in unsupplemented nests.The increasing amount of experimental data showing that brood culling is an important part of the biology of ants and other social insects emphasis the need to combine kin selection and life‐history approaches to study the outcome of queen–worker conflicts on the dynamics of colony growth and sex allocation.

Sociometry and sociogenesis of colonies of the harvester ant, Pogonomyrmex badius: distribution of workers, brood and seeds within the nest in relation to colony size and season

Summary1. The vertical distribution in the nest of chambers, workers, callow workers, brood and seeds was studied in the harvester ant, Pogonomyrmex badius, in northern Florida. On each of four sample dates (May, July, October, January), six to seven colonies, chosen to represent the full range of sizes, were excavated. All chamber contents were collected and counted. Chambers were mapped and measured. In a preliminary study, two nests were excavated after preventing vertical migration by driving barriers into the wall of a pit next to the nest, severing the vertical tunnels. The vertical distribution of these barrier‐nests differed little from unrestrained nests, indicating that unrestrained excavation produced a reasonable picture of vertical distributions.2. Nest depth, chamber number and total area increased with colony size. Chamber area declined sharply with depth, as did chamber number, such that more than half of the total area was found in the upper quarter of the nest.3. The proportion of dark‐coloured (older) workers also declined strongly with depth, but this decline was weaker in the spring, and depended to a modest degree on colony size. Conversely, in the distribution of callow (young) workers, the proportion increased towards the bottom of the nest. Mean worker age was inversely related to the depth at which workers were found. The proportion of the brood also increased towards the bottom of the nest, with worker brood, sexual brood, pupae and larvae all being distributed similarly.4. By contrast, seeds were stored at a preferred absolute depth between 40 and 100 cm. Colonies shallower than 100 cm stored seeds in their deepest chambers. Larger colonies stored most seeds in the upper third of the nest, but patterns were somewhat erratic because chambers were either filled completely with seeds or were empty.5. Because chamber area decreased sharply with depth, the densities (individuals cm–2) of all colony members, including dark workers, were lowest near the surface and highest in the deepest parts of the nest. Here, worker densities ranged from 2 to 8 cm–2, and brood from 2 to 25 cm–2.6. The regularity of the patterns of distribution suggests that harvester ant colonies have considerable spatial and temporal structure, which serves or is the outcome of important colony processes. A simple mechanism that could generate several of these patterns is discussed. New workers produced deep in the nest move upwards as they age. As they leave the brood zone they change from brood care to general nestduties, including increased nest excavation, leading to the top‐heavy pattern of nest area. As they appear at the surface, they change to guarding and foraging. Thus, age polyethism may be partly the result of this upward migration of workers.

Field Studies of the Parasitic AntSolenopsis daguerrei(Hymenoptera: Formicidae) on Fire Ants in Argentina

The workerless parasitic ant Solenopsis daguerrei (Santschi) has been considered a potential candidate for biological control of imported fire ants in the United States since the 1970s. We estimated the detrimental effects of S. daguerrei in fire ant colonies from Argentina by comparing the composition of parasitized and nonparasitized field colonies of the black imported fire ant S. richteri Forel. We recorded mound density and volume, incidence of polygyny, number of queens per colony, queen weight, and presence and abundance of brood and sexuals. Also, we studied aspects of the parasite’s biology such as parasitization rates, presence of sexuals, abundance and weight of queens, and sex ratio. S. daguerrei occurred in 1.2–23.7% of the colonies at 21 collecting sites. The mound density at parasitized sites was significantly lower than in parasite-free sites (161 versus 239 mounds per hectare). Parasitized colonies had significantly fewer host queens than nonparasitized ones (2.9 versus 5.5 queens per colony). The percentage of colonies with worker brood was significantly lower (in fall) in parasitized than in nonparasitized colonies. Also, worker brood was significantly less abundant (in fall and spring) in parasitized colonies. A short delay in the production of sexual brood by the host was observed in parasitized colonies. These detrimental effects look promising but should be evaluated in the long-term and confirmed in other areas. A better understanding of the host and parasite is required to introduce S. daguerrei into the United States to control imported fire ants.

Relationship Between Colony Size of Solenopsis richteri (Hymenoptera: Formicidae) and Infection with Thelohania solenopsae (Microsporida: Thelohaniidae) in Argentina

We studied the association between the size and composition of field colonies of the black imported fire ant, Solenopsis richteri Forel, and the presence of the microsporidian disease Thclohania solenopsae Knell, Allen,& Hazard in Buenos Aires Province, Argentina. We compared volumes of infected colonies with those of healthy colonies within an area where infected colonies were common and with those of healthy colonies from an area free of the infection. We also compared presence of polygyny, numbers of queens, presence and abundance of sexuals, broods, and myrmecophiles. Mound volume of infected colonies was substantially smaller than that of healthy colonies from the same area and from the disease-free area (4.9 versus 14.7 and 18.7 liters, respectively). Polygyny, number of queens per colony, presence of worker broods, and presence of winged females and males did not differ significantly in infected and healthy colonies. Sexual broods were less frequent in infected than in healthy colonies (17 versus 37%, respectively). Presence of myrmecophiles was as frequent in infected as in healthy colonies, but tile bug Blissus parasigaster (Bergroth) was more numerous in infected colonies. T. solenopsae may be a good candidate for the biological control of imported fire ants in the United States.

Elimination of sexual brood in the Argentine ant Linepithema humile: queen effect and brood recognition

AbstractMated queens of the ant Linepithema humile (Iridomyrmex humilis Mayr) introduced into dequeened colony fragments rearing sexual brood elicited worker aggression resulting in queen larvae being bitten and eliminated. By contrast, male larvae were spared. Regarding queen brood, killing mainly concerned small and medium sized larvae. A large proportion of the large larvae escaped extermination, and prepupae and pupae were spared. These data suggest that workers were able to discriminate sex, caste and age of the brood.That a queen pheromone may be involved was shown by experiments using whole or cut corpses that were either rinsed or not rinsed in pentane. The pheromone eliciting worker aggressive behaviour was shown to act over a short distance, suggesting that it is somewhat volatile. Similarities and differences between this new queen pheromone and other known queen pheromones acting on queen production or worker attraction are discussed as well as the origin of the signals underlying the recognition of the larval classes.