Reproductive allocation in multinest colonies of the ponerine ant Pachycondyla goeldii

Abstract.  1.  Pachycondyla goeldii constitutes the only recorded case of a monogynous (i.e. one queen per colony) polydomous (i.e. several nests per colony) species in the Ponerinae subfamily. This study examines the impact of polydomy on reproductive allocation between nests (also called 'calies' in polydomous society) in Pachycondyla goeldii Forel, by: (i) recording the number of workers and sexuals in 67 nests belonging to 21 colonies; (ii) dissection of workers in nine nests containing a queen (QR nests), nine nests without a queen but associated to a QR nest (QL nests) and five nests belonging to colonies that permanently lost the queen (OR nests); and (iii) measuring the length of all eggs present in the nests (our laboratory study shows that queen-laid eggs were significantly longer than worker-laid eggs).
2. The number of workers was significantly higher in QR nests than in QL nests, while the number of virgin queens was significantly higher in QL nests compared with QR nests.
3. Worker ovarian activity is inversely related to queen proximity: highest in OR nests, intermediate in QL nests, and lowest in QR nests.
4. Egg length was highest in QR nests, where the queen was most likely the primary egg-layer, intermediate in QL nests, where eggs could have originated from both the queen and workers, and lowest in OR nests, where workers were the sole egg-layers.
5. We postulate that the proximal mechanism explaining differences between QR and QL nests is the pheromonal absence of the queen from QL nests and that the evolutionary reasons of these divergences between nest types are likely to originate from the different conflicts occurring in ant colonies.     

Nest demographics and foraging behavior of <Emphasis Type="Italic">Apterostigma collare</Emphasis> Emery (Hymenoptera,Formicidae) provide evidence of colony independence

Apterostigma collare Emery is a highly derived fungus-growing ant within the Tribe Attini whose small, fungal nests are found in tropical rain forests. This study focuses on determining the colony structure of A. collare, specifically searching for evidence of polydomy or independence. We surveyed and observed nests in the field, and performed foraging bioassays and dissected nests in the laboratory. We determined the size and contents of nests in field populations. Nests found near other nests were not statistically different in size compared to nests found alone. There was also no statistical difference between near and lone nests regarding the presence of a queen in the nest. Most nests contained one queen with brood and workers, regardless of their proximity to other nests. Observations also were made of foraging and trail-marking behaviors. Foraging activity observed in the field revealed that workers left the nest area and followed trails upwards into the canopy, but they did not interact with foragers from other nearby nests. In a laboratory foraging arena, foragers marked a trail to a food source by dragging the gaster. Bioassays showed that A. collare workers preferred their own foraging trails, but not those of other conspecific colonies. All results suggest that each nest represents an independent colony, supporting a previous report that nests found in close proximity do not constitute a polydomous colony. Received 19 July 2006; revised 23 March 2007; accepted 6 June 2007.     

Reproductive allocation within a polygyne, polydomous colony of the ant Myrmica rubra

1. Ant colonies commonly have multiple egg‐laying queens (secondary polygyny). Polygyny is frequently associated with polydomy (single colonies occupy multiple nest sites) and restricted dispersal of females. The production dynamics and reproductive allocation patterns within a population comprising one polygyne, polydomous colony of the red ant Myrmica rubra were studied. 2. Queen number per nest increased with nest density and the number of adult workers increased with the number of resident queens and with nest density. This suggests that nest site limitation promotes polygyny and that workers accumulate in nest units incapable of budding. 3. Nest productivity increased with the number of adult workers and production per queen was independent of queen number. Productivity increased with nest density, suggesting local resource enhancement. This shows that productivity can be a linear function of queen numbers and that the limiting factor is not the egg‐laying capacity of queens. 4. The total and per capita production of reproductives decreased towards the periphery of the colony, suggesting that the spatial location of nest units affects sexual production. Thus nests at the periphery of the colony invested more heavily in new workers. This is consistent with earlier observations in plants and could either represent investment in future budding or increased defence. 5. The colony produced only five new queens and 2071 males, hence the sex ratio was extremely male biased.     

Seasonal spatial dynamics and causes of nest movement in colonies of the invasive Argentine ant (Linepithema humile)

Abstract.  1. Colony organisation and movement behaviour of the Argentine ant ( Linepithema humile ) was studied over 3 years in field populations in California and in captive colonies in the laboratory. This invasive species is highly polydomous and unicolonial; colonies consist of expansive and fluid networks of nests and trails. The spatial and temporal organisation of colonies may contribute to ecological dominance.
2. Argentine ant nests and inter-nest trails shift in size, abundance, and location, so that colony networks are spatially contracted in the winter and expanded spring to autumn. Colonies occupy permanent sites; ants migrated to and from the same winter nest locations year after year, and occupied 30% of the same nests repeatedly during seasonal migrations.
3. Nests were moved on average 2–3 m. Forty-two per cent were occupied less than 1 month, 4% the entire study, and the other 54% lasted 3.9 ± 2.3 months (mean ± SD).
4. Nests were located within 2–4 m of woody plants, in warm sites in the winter and cool sites in the summer. Both humidity and food availability influenced nest-site choice in laboratory colonies. However, when faced with a trade-off between factors, the ants chose humid nest boxes over nest boxes near food, and ants moved nests only in response to changes in humidity and not distance to food.
5. The results indicate that L. humile colonies are seasonally polydomous, and that nest movements are driven by changes in microclimate. Colony organisation maintains high local density and increases food supply, which may improve the competitive ability of L. humile colonies and reduce opportunities for species coexistence.     

Suburban sprawl: environmental features affect colony social and spatial structure in the black carpenter ant,Camponotus pennsylvanicus

1. In social insects, the number of nests that a colony inhabits may have important consequences for colony genetic structure, the number of queens, sex allocation, foraging efficiency, and nestmate recognition. Within the ants, colonies may either occupy a single nest (monodomy) or may be organised into a complex network of nests and trails, a condition known as polydomy. 2. The current study is a large‐scale, long‐term, comprehensive field examination of various features of colony social and spatial structure in the facultatively polydomous black carpenter ant, Camponotus pennsylvanicus (DeGeer). The study examined the density, persistence, and the spatiotemporal distribution of colonies across a gradient of land disturbance associated with urban development. The temporal and spatial pattern of nest use was compared between fragmented landscapes where nesting sites were interspersed among human‐built structures (urban plots) and less disturbed landscapes with higher tree density (suburban plots). In addition, nesting site fidelity and changes in colony spatial structure were monitored over 7 years. 3. Long‐term monitoring and extensive sampling over a large spatial area allowed the first comprehensive insight into the spatiotemporal dynamics of colony and population structure in C. pennsylvanicus. A total of 1113 trees were inspected over 233 ha. Camponotus pennsylvanicus were active on 348 of the 1113 trees (31%) and these represented 182 distinct colonies. The colonisation rate remained relatively stable over 7 years suggesting that an equilibrium point had been reached. Relative to the suburban plots, tree density was 65% lower in the urban plots. The proportion of trees colonised by C. pennsylvanicus was significantly higher in the urban plots suggesting that intraspecific competition for nesting sites may be especially high in areas with lower tree density. Colony spatial structure also differed significantly between habitats and a higher incidence of monodomy was observed in the urban environment. The average number of trees per colony across all subplots was 1.95 (range 1–4) indicating that C. pennsylvanicus are weakly polydomous. 4. The composite picture that emerges for C. pennsylvanicus colonies in the urban habitat is a chain reaction of events: (i) the urban habitat has a lower tree density, (ii) lower tree density results in higher tree colonisation rate, (iii) higher tree colonisation rate results in simpler colony spatial structure (i.e. higher incidence of monodomy), and (iv) simpler colony spatial structure results in numerically smaller colonies. Long‐term monitoring of the spatiotemporal pattern of nest site use in selected colonies revealed a unique trend. While worker counts in selected colonies remained relatively stable throughout the course of the study, colony spatial structure changed considerably with 28% of colonies experiencing a change. Furthermore, the likelihood of detecting a change in colony spatial structure increased with the amount of time passing from the initial inspection. 5. In conclusion, tree density has a significant effect on a number of important colony features in C. pennsylvanicus. Besides tree density, other environmental features such as human‐built structures cause habitat fragmentation and may act as natural barriers to worker dispersal and/or foraging. Such barriers may ultimately affect the social and/or spatial structure at both the colony and the population level.     

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.     

Polydomy and the organization of foraging in a colony of the Malaysian giant ant Camponotus gigas (Hym. / Form.)

In Kinabalu National Park, Borneo we observed four colonies of the Malaysian giant ant Camponotus gigas in a primary forest. These predominantly nocturnal ants have underground nests, but forage in huge three-dimensional territories in the rain forest canopies. The colony on which our study was mainly focused had 17 nests with about 7000 foragers and occupied a territory of 0.8 ha. To improve observation and manipulation possibilities, these nests were linked at ground level by 430 m of artificial bamboo trail. A group of specialist transport worker ants carried food from `source' nests at the periphery to the central `sink' nest of the queen. Transport of food between nests started immediately after the evening exodus of the foragers. Transporter ants formed a physical subcaste among the minors and behaved according to predictions of the central-place foraging theory. Their load size was about five times that of the average forager and grew proportionally with head width. Longer distances were run by ants with greater head width and larger gross weight. Transporter ants that ran more often took heavier loads. Experiments with extra-large baits revealed that C. gigas used long-range recruitment to bring foragers from different nests to “bonanzas” at far distant places. The foraging strategy of C. gigas is based on a polydomous colony structure in combination with efficient communication, ergonomic optimization, polyethism and an effective recruitment system. Received: 16 March 1998 / Accepted: 24 August 1998     

Brood raiding and the population dynamics of founding and incipient colonies of the fire ant, Solenopsis invicta

Abstract. 1. One of the first activities of minim workers in incipient fire ant nests is mutual brood raiding, the amalgamation of nests through the reciprocal stealing of brood and defection of workers.
2. Discrete mating flights created cohorts of incipient colonies. About 25% of founding nests survived the claustral period of 16–48 days (depending on season). Early incipient colony mortality was 5–6% per day. Over 60% of this mortality was accounted for through brood raiding. Most colonies raided when each cohort first became active, and many raided more than once. Raid size (number of nests, duration, distance) tended to increase during the summer as colonies grew through raiding. After the raiding period, nest mortality rate dropped 3–10-fold. Only 1–3% of founding nests were still alive at this time.
3. Queens from losing or failed nests tended to abandon their nests and attempted to enter successful ones, often following raiding trails to do so. This emigration was at least as successful as non-emigration in ultimately achieving the status of reproductrix of a successful nest (about 4%).
4. Brood raiding is a dominant process in early population dynamics, probably accounting for most of the early nest mortality. Its effect is to change the venue and unit of competition from nest-against-nest to a shifting aggregation of queens, workers and brood involving entire local populations. Nest thinning is thus very rapid, and the boost to the size of winning nests very large, allowing raiding colonies to win the competition for territory, and to achieve the early colony maturity so important to this, and other, weedy species. The importance of winning brood raids may also have driven increased minim production through the evolution of pleometrosis.     

Colony characteristics influence the risk of nest predation of a polydomous ant by a monotreme

Although predation of individual social insect workers has little effect on colony fitness, nest predation may be a significant selective agent because it can result in substantial loss of reproductive success. Surprisingly, the consequences of predation on social insect nests are poorly understood. In the present study, we investigate the factors that correlate with the probability of predation by echidnas, Tachyglossus aculeatus , on nests and colonies of the facultatively polydomous meat ant, Iridomyrmex purpureus. In particular, we investigate whether colony fragmentation provides a mechanism for reducing the costs of echidna predation. Over 2 years, 138 of the 140 colonies on our study site were depredated. Nest predation was most common in woodlands but with no obvious seasonal patterns. The probability of nest predation was positively correlated with the size of the nest, and negatively correlated with the density of surrounding nests. Although polydomous colonies are at a similar risk of predation by echidnas, the proportion of depredated nests is negatively correlated with the number of nests; thus, the probability that one or more nests avoid predation is increased with increasing nest numbers. Surprisingly, we found no influence of the level of echidna predation on colony growth, measured by either changes in the number of nests or the number of nest entrance holes.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 1–8.     

Colony productivity and foundress behaviour of a native wasp versus an invasive social wasp

Abstract.  1. Colony productivity, prey utilisation, and foundress behaviour of a North American native wasp ( Polistes fuscatus ) versus an European invasive wasp ( Polistes dominulus ) were investigated in a controlled field experiment with optimal versus natural foraging conditions. Colonies with the optimal prey foraging conditions were provided with prey ad libitum within an enclosed area. The other colonies foraged in the adjacent field–woodland but had the same nest conditions as the other treatment.
2. When given prey ad libitum , both wasp species captured similar amounts of prey and the conversion to total offspring biomass was similar. But P. dominulus colonies produced 2.5 times the number of workers as P. fuscatus colonies, reflecting the smaller size of P. dominulus wasps.
3. Foundresses of P. dominulus were observed more often building or repairing the nest, thereby contributing to the production of colonies with twice as many cells as colonies of P. fuscatus . Foundresses of P. dominulus showed more acts of aggression toward workers than did P. fuscatus foundresses, which was not a function of adult density on the nest.
4. At the end of the experiment, P. dominulus colonies with optimal prey foraging conditions still had a high level of egg-laying and peaked in the number of pupae then, whereas egg-laying and the number of pupae per colony of the other treatments began to decline 2–3 weeks earlier. These results indicate that P. dominulus is more opportunistic than P. fuscatus , which may account in part for P. dominulus 's success as an introduced species in North America.     

Pirate ants (Polyergus breviceps) and sympatric hosts (Formica occulta and Formica sp. cf. argentea): host specificity and coevolutionary dynamics

The pace and trajectory of coevolutionary arms races between parasites and their hosts are strongly influenced by the number of interacting species. In environments where a parasite has access to more than one host species, the parasite population may become divided in preference for a particular host. In the present study, we show that individual colonies of the pirate ant Polyergus breviceps differ in host preference during raiding, with each colony specializing on only one of two available Formica host species. Moreover, through genetic analyses, we show that the two hosts differ in their colony genetic structure. Formica occulta colonies were monogynous, whereas Formica  sp. cf. argentea colonies were polygynous and polydomous (colonies occupy multiple nest sites). This difference has important implications for coevolutionary dynamics in this system because raids against individual nests of polydomous colonies have less impact on overall host colony fitness than do attacks on intact colonies. We also used primers that we designed for four microsatellite loci isolated from P. breviceps to verify that colonies of this species, like other pirate ants, are comprised of simple families headed by one singly mated queen.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 565–572.     

Colony structure and spatial distribution of food resources in the polydomous meat ant <Emphasis Type="Italic">Iridomyrmex purpureus</Emphasis>

Polydomous social insects may reduce the costs of foraging by the strategic distribution of nests throughout their territory or home-range. This efficiency may most likely be achieved if the resources are relatively stable in place and time, and the colonies and nests are distributed in response to the location of the resources. However, no study has investigated how the distribution of food sources influences the spatial patterns of nests within polydomous colonies under natural conditions. Our two year study of 140 colonies of the Australian ant Iridomyrmex purpureus revealed that the decentralization of nests within colonies is associated with the distribution of trees containing honey-dew producing hemiptera. We show there is a positive correlation between the maximum distance between trees containing hemiptera and the maximum distance between nests within a colony. In addition, we demonstrate the mechanism by which this pattern may arise; new nests are built nearer to trees containing hemiptera than existing nests. Further, the distance between trees containing hemiptera and the nearest nests was negatively correlated with the length of exploitation of that tree. Finally, we show that most food is delivered to the nearest nest after which other ants redistribute it between the nests. Combined, these data suggest that foraging efficiency may be an important selection pressure favouring polydomy in I. purpureus. Received 6 April 2006; revised 29 September; accepted 4 October 2006.     

Colony structure, seasonality and food requirements of the crazy ant, Anoplolepis longipes (Jerd.), in the Seychelles

Abstract. 1. In Seychelles, A. longipes nests on the ground and in trees, underground nests do not occur, possibly because soils are too hard or unstable.
2. Nest density approached 700 ha^-1 in some areas and nests contained, on average, about 4000 ants. The total population (including foragers) in heavily infested areas may exceed 10 millions ha^-1.
3. On average, each nest contained about forty queens and fifty males, although only 50% of nests contained males. A few nests contained up to 300 queens, 1000 males, 36 000 workers and 23 000 brood.
4. Production of sexual adults and brood was related to the wet season, most being produced shortly before or after the heavy rains from November to March.
5. Natural spread is probably by budding of colonies which, during an expansive period, may advance 1 m day^-1.
6. Food collected by the colony included sugary substances from fruits, honey-dew-producing insects and plant exudates, and proteinaceous material, mainly in the form of insect prey and carcases. A forager collected, on average, 1.2 mg of liquid or 2.8 mg of solid material on each foraging trip.
7. Foraging occurred at fluctuating intensities throughout the day and night, but was limited by heavy rain, strong winds and substrate temperatures above 30°C. Maximum activity occurred at ground temperatures of 25–30°C and ceased about 44°C.
8. Availability of food and nest sites proably have the greatest influence on population size.     

The queen colony phase of vespine wasps (Hymenoptera, Vespidae)

Characteristics of queen nests of the Vespinae are reviewed including the number of cells and each brood stage in the mature nest, cell-building and egg-laying rates, length of each brood stage and the occurrence of brood cannibalism. Success rates of queen nests of the Vespinae are reviewed with their causes of failure including the effect of queen usurpation, and with particular reference to variation of queen behaviour or queen quality. Evidence for the differences between potentially successful and unsuccessful queen nests is given. Simulation models of queen nests are developed for potential successful and unsuccessful colonies to determine the extent of larval cannibalism, larval life extension and for unsuccessful queen nests, oophagy. The variation of cell building rate, the importance of ambient temperature and the fast development of the first batch of eggs are considered in relation to polistine studies. The causes of the variation of queen quality are unknown but could be related to the amount of food received as larvae or as adults before leaving the natal colony.     

Spatial foraging patterns and colony energy status in the African honey bee,Apis mellifera scutellata

The relationship between changes in foraging patterns (inferred from waggle dance activity) and colony energy status (inferred from brood rearing activity, food storage, and colony weight) was examined for the African honey bee during a period of relative resource abundance and resource dearth. When resources were more abundant mean foraging distances (about 400 m) and foraging areas (4–5 km²) were small, and colonies recruited to 12–19 different sites per day. Colony foraging ranges and sites visited increased slightly during the dearth period, yet foraging continued to be concentrated within less than 10 km². The degree to which fluctuations in foraging patterns were correlated with colony energy status varied with the availability of floral resources. During periods of relative forage abundance, increases in foraging range and number of sites visited were significantly correlated with increases in brood rearing and colony weight. In contrast, colonies examined during periods of resource dearth exhibited no correlations between foraging areas, foraging distances, and fluctuations in brood rearing, food storage, or colony weight. Thus, during dearth periods colonies may not be able to coordinate foraging patterns with changes in colony energy status.     

Foraging decisions of individual workers vary with colony size in the greenhead ant Rhytidoponera metallica (Formicidae, Ectatomminae)

Summary. The ability of worker ants to adapt their behaviour depending on the social environment of the colony is imperative for colony growth and survival. In this study we use the greenhead ant Rhytidoponera metallica to test for a relationship between colony size and foraging behaviour. We controlled for possible confounding ontogenetic and age effects by splitting large colonies into small and large colony fragments. Large and small colonies differed in worker number but not worker relatedness or worker/brood ratios. Differences in foraging activity were tested in the context of single foraging cycles with and without the opportunity to retrieve food. We found that workers from large colonies foraged for longer distances and spent more time outside the nest than foragers from small colonies. However, foragers from large and small colonies retrieved the first prey item they contacted, irrespective of prey size. Our results show that in R. metallica, foraging decisions made outside the nest by individual workers are related to the size of their colony.Received 23 March 2004; revised 3 June 2004; accepted 4 June 2004.     

Serial monodomy in ants: an antipredator strategy?

Abstract.  1. The term serial monodomy is used to describe a life-history phenomenon in social insects. Serially monodomous colonies maintain multiple nests for their exclusive use, but only occupy one nest at a time.
2. The hypothesis that colony odours mediate nest relocation decisions was tested in the serially monodomous species Aphaenogaster araneoides from Central America. Odour extracts of colony members were created using a non-polar solvent.
3. Colonies strongly avoided reoccupying nests treated with colony odour extracts, while control colonies often returned to nests subjected to solvent-only control. Behavioural observations indicated that A. araneoides colonies are capable of detecting army ant ( Eciton burchellii ) raids up to 1.4 m from the raiding front, with several seconds to evacuate nests.
4. It is proposed that the function of serial monodomy in A. araneoides is the reduction of nest odour to enhance detection of predaceous army ants. Serial monodomy may be a widespread but undocumented mode of nesting where army ants occur in tropical and subtropical climates.     

Colony Budding and its Effects on Food Allocation in the Highly Polygynous Ant, Monomorium pharaonis

To advance our understanding of the causes and the consequences of budding (colony multiplication by fragmentation of main nests), we investigated nest movement in the facultatively polydomous Pharaoh ant, Monomorium pharaonis. Demographic data revealed that Pharaoh ants are highly polygynous and have a relatively low worker to queen ratio of 12.86. Budding experiments demonstrated that the number of available bud nests has a significant effect on colony fragmentation and increasing the number of bud nests resulted in smaller colony fragments. The overall distribution among bud nests was uneven, even though there was no evidence that the different life stages and castes partitioned unevenly among the bud nests and the analysis of individual colonies revealed no evidence of an uneven split in any of the colonies. This demonstrates that Pharaoh ants have the ability to exert social control over colony size and caste proportions during budding, which may contribute to their success as an invasive ant. The intensity of nest disturbance had a significant effect on whether or not the ants migrated into bud nests. Major disturbance resulted in the ants abandoning the source nest and migrating to bud nests and minor disturbance did not stimulate the ants to abandon the source nest. The results of the successive budding experiment which allowed the ants the opportunity to bud into progressively smaller nest fragments demonstrate that Pharaoh ants maintain a preferred minimum group size of 469 ± 28 individuals. Food allocation experiments utilizing protein marking revealed that nest fragmentation in Pharaoh ants has no negative impact on intracolony food distribution. Overall, our results suggest that nest units in the Pharaoh ant behave like cooperative, rather than competitive, entities. Such cooperation is most likely facilitated by the fact that individuals in all bud nests are genetically related, remain in close proximity to each other, and may continue to exchange individuals after budding.     

Microgeographic genetic structure and intraspecific parasitism in the ant Leptothorax nylanderi

1. Genetic colony structure of the small central European ant Leptothorax nylanderi is affected strongly by ecological constraints such as nest site availability and intraspecific social parasitism. 2. Although L. nylanderi is generally monogynous and monandrous, more than a quarter of all nests collected in a dense population near Würzburg, Germany, contained several matrilines. As shown by microsatellite analysis, the average nest‐mate relatedness in these nests was 0.20. Genetically heterogeneous nests arise from nest take‐over by alien colonies or founding queens, a result of severe competition for nest sites. 3. In summer, more than one‐third of all colonies inhabited several nest sites at a time. Polydomy appears to be rather limited, with two or three nests belonging to a single polydomous colony. 4. Queens appear to dominate male production; only a small fraction (8%) of males was definitively not progeny of the queen present but might have been worker progeny or offspring of another queen. 5. Strong evidence for heterozygote deficiency was found and a total of nine diploid males was discovered in two colonies. These findings suggest deviation from random mating through small, localised nuptial flights.     

Resource characteristics and competition affect colony and individual foraging strategies of the wood ant Formica integroides

Abstract.  1. Resource characteristics and competitive pressure can affect an ant colony's foraging strategy. This study examined the ability of the wood ant Formica integroides to respond, at both the colony and individual levels, to changes in competitive pressure for access to terrestrial and arboreal resources.
2. Because foraging behaviours depend on resource characteristics, foraging for different resource types (e.g. terrestrial and arboreal habitats) produces different spatial or territorial arrangements. In this study, terrestrial contests for resources followed an interference-exploitation tradeoff, while arboreal foragers defended entire trees as absolute territories.
3. Competitive pressure for access to arboreal resources was shown to increase with distance from F. integroides nests.
4. In this study, the ability of F. integroides to defend a resource varied with body size. Large foragers were better defenders than small foragers. For groups of foragers, the ability to defend a resource increased with the ratio of large to small foragers.
5. In response to competitive pressure, F. integroides colonies altered the size distribution of arboreal, but not terrestrial, foragers. An increase in competitive pressure was matched by an increase in the number of large foragers allocated to trees. This response to competition affected the relationship between body size and distance from the nest for arboreal foragers.
6. Foraging behaviours for individual arboreal foragers also varied with competitive pressure. As competition increased, large arboreal foragers spent more time in direct contact with the resource rather than standing between resource patches.