Nest Enlargement in Leaf-Cutting Ants: Relocated Brood and Fungus Trigger the Excavation of New Chambers

During colony growth, leaf-cutting ants enlarge their nests by excavating tunnels and chambers housing their fungus gardens and brood. Workers are expected to excavate new nest chambers at locations across the soil profile that offer suitable environmental conditions for brood and fungus rearing. It is an open question whether new chambers are excavated in advance, or will emerge around brood or fungus initially relocated to a suitable site in a previously-excavated tunnel. In the laboratory, we investigated the mechanisms underlying the excavation of new nest chambers in the leaf-cutting ant Acromyrmex lundi. Specifically, we asked whether workers relocate brood and fungus to suitable nest locations, and to what extent the relocated items trigger the excavation of a nest chamber and influence its shape. When brood and fungus were exposed to unfavorable environmental conditions, either low temperatures or low humidity, both were relocated, but ants clearly preferred to relocate the brood first. Workers relocated fungus to places containing brood, demonstrating that subsequent fungus relocation spatially follows the brood deposition. In addition, more ants aggregated at sites containing brood. When presented with a choice between two otherwise identical digging sites, but one containing brood, ants'' excavation activity was higher at this site, and the shape of the excavated cavity was more rounded and chamber-like. The presence of fungus also led to the excavation of rounder shapes, with higher excavation activity at the site that also contained brood. We argue that during colony growth, workers preferentially relocate brood to suitable locations along a tunnel, and that relocated brood spatially guides fungus relocation and leads to increased digging activity around them. We suggest that nest chambers are not excavated in advance, but emerge through a self-organized process resulting from the aggregation of workers and their density-dependent digging behavior around the relocated brood and fungus.     

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

1. 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 nest duties, 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.     

Nest architecture of the ant <Emphasis Type="Italic">Formica pallidefulva</Emphasis>: structure,costs and rules of excavation

Summary The architecture of underground ant nests was studied in the ant Formica pallidefulva. These ants build shallow (30–45 cm deep) nests, which consist of more or less vertical shafts that bear chambers. Shafts are modular units of nest growth; nests are enlarged by adding more shafts or extending previously existing ones. The nests are top-heavy, their volume declining exponentially with depth. The total volume of the nest is strongly correlated with the number of worker occupying the nest (R² = 0.87). Some of the rules and templates used by workers for nest construction were determined: (a) chambers are formed in the direction of the tunnels leading up to them, (b) the amount of soil excavated per unit time increases with soil temperature and moisture content. The amount of time and energy required to construct a typical nest were approximated using digging ability parameters determined in the lab. We estimate that if a colony was to move twice a year, it would expend around 20% of its energy intake and at least 6% of its worker time on nest excavation.Received 16 December 2002; revised 10 July 2003; accepted 23 July 2003.     

Life Inside an Acorn: How Microclimate and Microbes Influence Nest Organization in Temnothorax Ants

Nests provide a buffer against environmental variation, but conditions may also vary at different locations within a nest. Conditions can vary based on abiotic factors, such as moisture and temperature, as well as biotic factors, such as the presence of microbes and potential pathogens. Therefore, characterizing how animals adjust their position inside their nests to track microclimate preferences while at the same time preventing pathogen exposure is necessary to understand the benefits nests provide. Here we studied how colonies of the acorn‐nesting ant Temnothorax curvispinosus responded to experimental manipulation of moisture, temperature, and microbial growth inside their nests. Colonies showed no response to differences in moisture and moved to the bottom of the acorn regardless of moisture treatment. When nests were heated from the top to simulate warming by the sun, workers preferentially moved brood to the warm, upper half of the acorn, which would stimulate brood development. Finally, the strongest factor that influenced colony position was the presence of microbes inside the nest—colonies avoided the bottom of the nest when it was inoculated with microbes, and colonies in new acorns shifted to the top of the acorn over time as mold and other microbes had time to grow. The relatively strong response of T. curvispinosus to microbial growth inside their nests suggests that pathogen pressures—in addition to microclimate—have a significant impact on how colonies use nest spaces. Social insects are known to invest heavily in antimicrobial compounds that kill or slow the growth of microbes, but avoidance may represent an additional line of defense to prevent pathogen exposure.     

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

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

Avoiding predation by army ants: Defensive behaviours of three ant species of the genus Camponotus

Antipredation behaviours of three ant species of the genus Camponotus were investigated in field and laboratory studies. Contact with the army ant Neivamyrmex nigrescens was shown to produce nest evacuation and removal of brood by colonies of Camponotus festinatus. Colonies of Camponotus ocreatus and Camponotus vicinus defended their nests through recruitment of the major caste. The initiation of evacuation or aggressive recruitment was dependent upon tactile contacts between Camponotus nestmates. Other sorts of disturbance which did not involve army ant contact did not produce evacuation or recruitment, indicating a high degree of enemy specification by Camponotus. The comparison of worker morphology revealed a close relationship between each species' strategy for nest defence and the degree of worker polymorphism exhibited. These findings suggest that interspecific ant predation has been a significant factor in the evolution of colony defence systems and caste polymorphism.     

Ant Colonies Prefer Infected over Uninfected Nest Sites

During colony relocation, the selection of a new nest involves exploration and assessment of potential sites followed by colony movement on the basis of a collective decision making process. Hygiene and pathogen load of the potential nest sites are factors worker scouts might evaluate, given the high risk of epidemics in group-living animals. Choosing nest sites free of pathogens is hypothesized to be highly efficient in invasive ants as each of their introduced populations is often an open network of nests exchanging individuals (unicolonial) with frequent relocation into new nest sites and low genetic diversity, likely making these species particularly vulnerable to parasites and diseases. We investigated the nest site preference of the invasive pharaoh ant, Monomorium pharaonis, through binary choice tests between three nest types: nests containing dead nestmates overgrown with sporulating mycelium of the entomopathogenic fungus Metarhizium brunneum (infected nests), nests containing nestmates killed by freezing (uninfected nests), and empty nests. In contrast to the expectation pharaoh ant colonies preferentially (84%) moved into the infected nest when presented with the choice of an infected and an uninfected nest. The ants had an intermediate preference for empty nests. Pharaoh ants display an overall preference for infected nests during colony relocation. While we cannot rule out that the ants are actually manipulated by the pathogen, we propose that this preference might be an adaptive strategy by the host to “immunize” the colony against future exposure to the same pathogenic fungus.     

Nest structure and colony cycle of the Allegheny mound ant,Formica exsectoides Forel (Hymenoptera: Formicidae)

Summary The nest structure and colony cycle of a population of Allegheny mound ants,Formica exsectoides, were examined in central Michigan. The dispersion pattern of mounds was random. Nest structure and presence of brood were primarily determined by excavation of twenty-three nests over three intervals from June through September. Additional excavations of five nests in 1990 and ten nests in 1991 provided further details on nest structure and colony cycle. Most galleries occurred within the mound and upper 30 cm of soil, but some activity reached depths of 100 to 270 cm. Depth of nests showed little correlation with external measurements of height and diameter. Immature stages were recovered from two strata: the upper 20 cm of nest and mound and the lowest nest depths. Alate sexual forms were found in or near the mound in July, and numerous dealate queens were collected in September from peripheral galleries near the soil surface.     

Nest relocation in the ant Pheidole dentata

We studied nest relocation in the ant Pheidole dentata, a common species in the southern US, by following colonies for 6 weeks. We correlated probability of relocation with several abiotic and biotic environmental factors, such as air temperature, humidity, leaf litter depth (LLD), nest type and presence of phorid fly parasitoids. Colonies moved often, on average every 16.2 days. By the end of our study, fewer than 5 % of colonies marked at the start remained in their original nests. Only 3.1 % of colonies returned to a previously used nest. The mean distance of relocation was 0.76 ± 0.73 m. Colonies nested in a variety of locations, such as in the ground, by tree trunks, under rocks and inside wood. Several factors affected probability of relocation. Higher LLD and vapor pressure deficit (VPD) increased probability of relocation. Probability of relocation varied by nest type, by plot and week of study. We found no evidence that surface cover or rain affected relocation behavior. Colonies appear to avoid nests that become dry by nesting against tree trunks. Phorid fly abundance correlated negatively with VPD, yet it had no effect on colonies’ likelihood of relocation. We discuss other implications of frequent nest relocations.     

Carbon dioxide concentrations in the nests of the mud-dwelling mangrove ant Polyrhachis sokolova Forel (Hymenoptera: Formicidae)

Abstract The nests of the mangrove ant Polyrhachis sokolova are found in soil in intertidal mangrove communities, and are thus inundated at high tides for several hours. Some of the nest galleries are flooded, but others retain air pockets, to which the ants retreat. During and following inundation, we measured carbon dioxide concentrations in air samples collected from different levels in the nests and from artificial 'control' holes in the mud. To account for the relative contribution of different sources of carbon dioxide, we also measured the carbon dioxide production by individual ants (including larvae and pupae) and small samples of mud collected near the ant nests. Nest carbon dioxide concentrations were high (2.5−11%) during and immediately following inundation, but the concentrations in the upper regions of the nest fell as soil water levels receded. However, at depths>10 cm below the level soil surface, the carbon dioxide concentrations remained relatively high and stable (at approximately 2%) over the 11 days between one high tide and the next. The contribution of the mud (and associated microorganisms) to the carbon dioxide concentration in the nests was substantial, and the contribution of the respiration of the ants was approximately 10−15% of the total. The carbon dioxide concentrations in the nests of this species during high tides are among the highest recorded for insect nests, suggesting that these ants may have unusual physiological attributes to match the behavioural and ecological challenges associated with living in the intertidal zone.     

Biomantling and Bioturbation by Colonies of the Florida Harvester Ant,Pogonomyrmex badius

In much of the world, soil-nesting ants are among the leading agents of biomantling and bioturbation, depositing excavated soil on the surface or in underground chambers. Colonies of the Florida harvester ant, Pogonomyrmex badius excavate a new nest once a year on average, depositing 0.1 to 12 L (3 L average) of soil on the surface. Repeated surveys of a population of about 400 colonies yielded the frequency of moves (approximately once per year), the distance moved (mean 4 m), and the direction moved (random). The area of the soil disc correlated well with the volume and maximum depth of the nest, as determined by excavation and mapping of chambers. The population-wide frequency distribution of disc areas thus yielded the frequency distribution of nest volumes and maximum depths. For each surveyed colony, the volume of soil excavated from six specified depth ranges and deposited on the surface was estimated. These parameters were used in a simulation to estimate the amount of soil mantled over time by the observed population of P. badius colonies. Spread evenly, P. badius mantling would create a soil layer averaging 0.43 cm thick in a millennium, with 10–15% of the soil deriving from depths greater than 1 m. Biomantling by P. badius is discussed in the context of the ant community of which it is a part, and in relation to literature reports of ant biomantling.     

Worker populations of Formica lugubris Zett. nests in Irish plantation woods

Abstract. 1. Estimates of worker populations of Formica lugubris Zett. nests in Irish plantation woods were obtained using capture—mark—recapture methods. Forager numbers were estimated by paint-marking individuals on the foraging routes and 'recapturing' by traffic counts. Colony-size was estimated using Stradling's (1970) ³²P-radiolabelling technique. Worker brood was estimated by excavation and direct count.
2. Forager numbers ranged from 6906 to 64 686 (eleven nests), colony-size ranged from 9797 to 71 052 (five nests) and worker brood ranged from 9809 to 16 269 (four nests).
3. Forager number was highly correlated with the average traffic on individual routes and the nest forager populations declined rapidly in >30-year-old nests.     

Nest mounds of red wood ants (Formicaaquilonia): hot spots for litter-dwelling earthworms

A previously undocumented association between earthworms and red wood ants (Formicaaquilonia Yarr.) was found during an investigation of the influence of wood ants on the distribution and abundance of soil animals in boreal forest soil. Ant nest mounds and the surrounding soil of the ant territories were sampled. The ant nest mound surface (the uppermost 5-cm layer) harboured a much more abundant earthworm community than the surrounding soil; the biomass of the earthworms was about 7 times higher in the nests than in the soil. Dendrodrilusrubidus dominated the earthworm community in the nests, while in soils Dendrobaenaoctaedra was more abundant. Favorable temperature, moisture and pH (Ca content), together with abundant food supply (microbes and decomposing litter) are likely to make a nest mound a preferred habitat for earthworms, provided that they are not preyed upon by the ants. We also conducted laboratory experiments to study antipredation mechanisms of earthworms against ants. The experiments showed that earthworms do not escape predation by avoiding contact with ants in their nests. The earthworm mucus repelled the ants, suggesting a chemical defence against predation. Earthworms probably prevent the nest mounds from becoming overgrown by moulds and fungi, indicating possible mutualistic relationships between the earthworms and the ants. Received: 21 November 1996 / Accepted: 3 April 1997     

Ants under crowded conditions consume more energy

Social insects live in colonies consisting of many workers, where worker interactions play an important role in regulating colony activities. Workers interact within the social space of the nest; therefore, constraints on nest space may alter worker behaviour and affect colony activities and energetics. Here we show in the ant Temnothorax rugatulus that changes in nest space have a significant effect on colony energetics. Colonies with restricted nest space showed a 14.2 per cent increase in metabolic rate when compared with the same colonies in large uncrowded nests. Our study highlights the importance of social space and shows that constraints on social space can significantly affect colony behaviour and energy use in ants. We discuss the implications of our findings regarding social insects in general.     

CO2 efflux from subterranean nests of ant communities in a seasonal tropical forest,Thailand

Many ant species construct subterranean nests. The presence of their nests may explain soil respiration “hot spots”, an important factor in the high CO₂ efflux from tropical forests. However, no studies have directly measured CO₂ efflux from ant nests. We established 61 experimental plots containing 13 subterranean ant species to evaluate the CO₂ efflux from subterranean ant nests in a tropical seasonal forest, Thailand. We examined differences in nest CO₂ efflux among ant species. We determined the effects of environmental factors on nest CO₂ efflux and calculated an index of nest structure. The mean CO₂ efflux from nests was significantly higher than those from the surrounding soil in the wet and dry seasons. The CO₂ efflux was species‐specific, showing significant differences among the 13 ant species. The soil moisture content significantly affected nest CO₂ efflux, but there was no clear relationship between nest CO₂ efflux and nest soil temperature. The diameter of the nest entrance hole affected CO₂ efflux. However, there was no significant difference in CO₂ efflux rates between single‐hole and multiple‐hole nests. Our results suggest that in a tropical forest ecosystem the increase in CO₂ efflux from subterranean ant nests is caused by species‐specific activity of ants, the nest soil environment, and nest structure.     

Nest Relocation and Excavation in the Florida Harvester Ant,Pogonomyrmex badius

The Florida harvester ant (Pogonomyrmex badius) excavates deep nests in the sandy soils of the Gulf and Atlantic coastal plains. Nest relocations of over 400 colonies in a north Florida coastal plains pine forest were tracked and mapped from 2010 to 2013. Individual colonies varied from one move in two years to four times a year, averaging about one per year. Almost all moves occurred between May and November peaking in July when more than 1% of the colonies moved per day. Move directions were random, and averaged 4 m, with few moves exceeding 10 m. Distance moved was not related to colony size. Over multiple moves, paths were random walks around the original nest location. Relocation is probably intrinsic to the life history of this species, and the causes of relocation remain obscure— the architecture of old and new nests was very similar, and neither the forest canopy nor the density or size of neighbors was correlated with relocation. Monitoring entire relocations (n = 20) showed that they were usually completed in 4 to 6 days. Moves were diurnal, peaking in the mornings and afternoons dipping during mid-day, and ceasing before sundown. Workers excavated the new nest continuously during the daytime throughout the move and beyond. A minority of workers carried seeds, charcoal and brood, with seeds being by far the most common burden. The proportion of burdened workers increased throughout the move. Measured from year to year, small colonies gained size and large ones lost it. Colonies moving more than once in two years lost more size than those moving less often, suggesting that moving may bear a fitness cost. Colony relocation is a dramatic and consistent feature of the life history of the Florida harvester ant, inviting inquiry into its proximal and ultimate causes.     

Nest of the dinosaur Protoceratops

Thulborn, R A. 1992 04 15: Nest of the dinosaur Protoceratops. Lethaia , Vol. 25, pp. 145–149. Oslo. ISSN 0024–1164.
Two different types of nests and eggs have been attributed to Protoceratops , a primitive-looking ceratopsian dinosaur from the Upper Cretaceous of Mongolia. Comparisons with the reproductive traces of other Cretaceous dinosaurs reveal that conventional restorations of the Protoceratops nest are probably based upon the nest of an ornithopod dinosaur. Authentic nests of Protoceratops do exist, but these are smaller and less complex in structure than the nests of ornithopod dinosaurs. This misunderstanding, which has endured in popular and scientific literature for more than 50 years, underlies the widespread belief that Protoceratops laid its eggs in a crater-like excavation. More probably the nest of Protoceratops comprised a shallow radial array of eggs concealed beneath a low mound of soil. * Ornithischia, eggs, nests, behaviour, Cretaceous .     

Breeding behaviour, relatedness and sex-investment ratios inLeptothorax tuberum Fabricius

The ant speciesLeptothorax tuberum was shown to be predominantly monogynous. Queens usually mate once only but some nests may have a multiply-mated queen or are partially or serially polygynous. As expected from these results, within nest relatedness between workers and between workers and alate queens was found to be high. Almost fifty percent of nests had no nest queen which may indicate high queen mortality, queens leaving to found new nests or nest fragmentation. Observed female investment frequencies (IF _o ) were not significantly different from those expected on the basis of worker control of sexual production and the relatedness estimates of workers to alate queens and workers to males calculated from isozyme data (IF _E ). These values were not consistent with queen control. There was no evidence for lower IF _o s in queenless nests nor for higher IF _o s in larger nests classified by worker number. When nests were classified by sexual productivity, however, there was a strong rank correlation between productivity and female bias. This is the first study of an ant species to test observed IF against expected IF calculated without inferring between caste relatedness from worker data or pooling of data from different castes.     

Nest site selection by a neotropical swarm-founding wasp: seasonal alternation of nest orientation

Nest orientation in social insects has been intensively studied in warmer and cooler climates, particularly in the northern hemisphere. Previous studies have consistently shown that species subjected to these climatic conditions prefer to select mostly southern locations where the nests can gain direct sunlight. However, very little is known on nest orientation in tropical and subtropical social insects. We studied nest orientations initiated by swarms throughout a year in a Brazilian swarm-founding wasp, Polybia paulista von Ihering (Hymenoptera: Polistinae). Swarms selected various orientations as nest sites, but there was a particular trend in that swarms in the winter period (May–August) preferred to build northward-facing nests. This preference is opposite from that of social wasps observed in the northern hemisphere. Colonies of this species can potentially last for many years with continuous nesting, but nesting activities of colonies during the winter are severely limited due to cool temperature and a shortened day length. Northward-facing nests are warmer through the gain of direct solar heat during the winter period; consequently, choosing northward-facing sites may be advantageous for swarms in terms of a shortened brood development and shortened time needed to increase metabolic rates during warm-up for flight.     

Bait distribution among multiple colonies of Pharaoh ants (hymenoptera: Formicidae)

Pharaoh ant, Monomorium pharaonis (L.), infestations often consist of several colonies located at different nest sites. To achieve control, it is desirable to suppress or eliminate the populations of a majority of these colonies. We compared the trophallactic distribution and efficacy of two ant baits, with different modes of action, among groups of four colonies of Pharaoh ants. Baits contained either the metabolic-inhibiting active ingredient hydramethylnon or the insect growth regulator (IGR) pyriproxyfen. Within 3 wk, the hydramethylnon bait reduced worker and brood populations by at least 80%, and queen reductions ranged between 73 and 100%, when nests were in proximity (within 132 cm) to the bait source. However, these nest sites were reoccupied by ants from other colonies located further from the bait source. The pyriproxyfen bait was distributed more thoroughly to all nest locations with worker populations gradually declining by 73% at all nest sites after 8 wk. Average queen reductions ranged from 31 to 49% for all nest sites throughout the study. Even though some queens survived, brood reductions were rapid in the pyriproxyfen treatment, with reductions of 95% at all locations by week 3. Unlike the metabolic inhibitor, the IGR did not kill adult worker ants quickly, thus, more surviving worker ants were available to distribute the bait to all colonies located at different nest sites. Thus, from a single bait source, the slow-acting bait toxicant provided gradual, but long-term control, whereas the fast-acting bait toxicant provided rapid, localized control for a shorter duration.