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.     

Genetic analysis of colony structure in polydomous and polygynous ant populations

Most interactions between individuals of social insects occur in colonies. The correct identification of colonies is therefore crucial for most empirical studies which aim to test evolutionary theories based on properties at the colony level. In many ant populations, the identification of colonics is hampered by polydomy, i.e. that single colonics occupy several, spatially separated nests. Only few attempts have been made so far to develop genetical methods for analysing the structure of specific colonics. Three methods to solve this problem are presented: rare genotype sisterhoods (tracking rare genotypes or alleles), G -distance (a measure of genotypic heterogeneity derived from G -statistics), and neighbour relatcdness (estimates of genetic relatcdness for specific nest pairs). Our methods quantify the likelihood of nest pairs being con-colonial or non-colonial, and given sufficient genetical resolution, statistical tests can be applied. The methods proposed here arc applied to a highly polygynous population of the red ant, Myrmica sulcinodis. In this population single colonics are found to inhabit 1–4 nests, and both monodomous and polydomous colonies coexist in dense clusters of nests. This result is discussed with respect to the functional significance of polydomy. Further, the general application of the methods for determination of colony structure is discussed.     

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.     

Polydomy in red wood ants

Polydomy, a single colony spread between multiple nests, is a widespread life history strategy in ants. The mechanisms by which a polydomous colony functions, and the fitness benefits this nesting strategy provides, are poorly understood. Here we review what is known about polydomy in the well-studied and ecologically important Formica rufa group. We focus particularly on the ecological fitness benefits polydomy may provide to members of the F. rufa group. We discuss the well-documented association in this group between polygyny (multiple queens in a colony) and polydomy, and how this relationship may favour colony reproduction by budding. We argue that although polygyny and reproduction by budding may drive a colony to spread between multiple nests, the maintenance of prolonged communication between these nests needs further explanation in terms of fitness benefits. The potential benefits of polydomy in the F. rufa group are discussed, specifically how polydomy may help a colony: exploit resources, dominate spaces, or lower the cost of stochastic nest destruction. The potential consequences of polydomy for the social organisation of a colony are explored. We also highlight gaps in current knowledge, and suggest future research directions.     

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.     

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.     

Mechanisms of dispersed central-place foraging in polydomous colonies of the Argentine ant

Many species of ants occupy multiple nests, a condition known as polydomy. Because of their decentralized structure, polydomous colonies may be removed from some of the constraints associated with classic central-place foraging. We used laboratory and field experiments to assess the mechanisms involved in dispersed central-place foraging in polydomous colonies of the Argentine ant Linepithema humile, a widespread invasive species. Both in the laboratory and in the field, Argentine ants established new nests at sites located near food. Laboratory colonies of L. humile redistributed workers, brood and resources among nests in response to the spatial heterogeneity of food resources. In addition, laboratory colonies formed recruitment trails between nests in the context of foraging, providing a mechanism for the transport of material between nests. This highly flexible system of allocating nests, workers and brood throughout a colony's foraging area potentially increases foraging efficiency and competitive ability. The importance of polydomy as a determinant of competitive ability is underscored by its prevalence among ecologically dominant ants, including most, if not all, highly invasive species. Copyright 2000 The Association for the Study of Animal Behaviour.     

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.     

Nest mate recognition in ants with complex colonies: within- and between-population variation

The ability to recognize kin is widespread, and especially importantin highly social organisms. We studied kin recognition by assessingpatterns of aggression within and between nests of the antLeptothorax longispinosus. Colonies of this species can befractionated into subunits, a condition called polydomy. Theproblem of recognizing relatives is therefore more complexwhen those relatives can live in two or more different places.We hypothesized that spatial subdivision may have resulted ina stronger genetic component to kin recognition than in caseswhere colonies live in a single location. To test our hypothesiswe assessed recognition capabilities for two populations ofthis ant that differ in the complexity of their colonies. Ina New York, USA, population, polydomy is very common, and coloniesalso can have multiple queens. By contrast, a population inWest Virginia, USA, has colonies that typically are monogynousand rarely are polydomous. We conducted introductions of antsbetween different nests collected in the same neighborhood,with self-introductions and alien introductions as controls.Nests from the two populations showed corresponding differencesin their aggression towards intruders. For New York nests, the extent of genetic similarity was the single best predictor ofaggression, whereas for West Virginia nests aggression wasjointly influenced by genetic similarity and spatial distance.In both populations, we found nest pairs for which aggressionwas nonreciprocal; these probably reflect recognition errors by one of the nests. After the ants were maintained in the laboratoryfor 3 months, their aggression scores rose and fewer recognitionerrors were made. Thus nest-mate and colony-mate recognitionin this species are mediated primarily by endogenous cues (geneticsimilarity); the importance of exogenous cues for nest materecognition depends on the population's social system.     

First Record of Polydomy in a Monogynous Ponerine Ant: A Means to Allow Emigration Between Pachycondyla goeldii Nests

The ponerine ant Pachycondyla goeldii is a monogynous (i.e. one queen per colony) arboreal species that colonizes pioneer areas. Founding queens and first generation workers initiate their own ant garden by building a cardboard-like structure into which epiphyte seeds are integrated. Following the growth of the epiphyte, the colony establishes its nest within the root system. This particular nest-building behavior is crucial in an environment where suitable nest sites are rare. Nevertheless, the slow growing process of ant gardens does not allow this species to readily evacuate and find another refuge in the advent of an attack by a predator or worsening climatic conditions. Previous field studies of P. goeldii were performed after forest destruction and subsequent colonization by P. goeldii. As a result, the colonies studied where relatively young and monodomous (i.e. one nest per colony). Our study of nest composition, worker exchanges between ant gardens in the field, and dyadic encounters shows that mature colonies of P. goeldii are polydomous (i.e. multiple nests per colony). In ants, the association of polydomy with monogyny has infrequently been reported. To our knowledge, P. goeldii represents the first record of a Ponerinae exhibiting both these particular characteristics. Our field and laboratory experiments suggest that polydomy is adaptively advantageous in coping with the microclimatic instability of pioneer areas by providing colonies with easily accessible nests.     

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 .     

Oh sister,where art thou? Spatial population structure and the evolution of an altruistic defence trait

The evolution of parasite virulence and host defences is affected by population structure. This effect has been confirmed in studies focusing on large spatial scales, whereas the importance of local structure is not well understood. Slavemaking ants are social parasites that exploit workers of another species to rear their offspring. Enslaved workers of the host species Temnothorax longispinosus have been found to exhibit an effective post‐enslavement defence behaviour: enslaved workers were observed killing a large proportion of the parasites’ offspring. As enslaved workers do not reproduce, they gain no direct fitness benefit from this ‘rebellion’ behaviour. However, there may be an indirect benefit: neighbouring host nests that are related to ‘rebel’ nests can benefit from a reduced raiding pressure, as a result of the reduction in parasite nest size due to the enslaved workers’ killing behaviour. We use a simple mathematical model to examine whether the small‐scale population structure of the host species could explain the evolution of this potentially altruistic defence trait against slavemaking ants. We find that this is the case if enslaved host workers are related to nearby host nests. In a population genetic study, we confirm that enslaved workers are, indeed, more closely related to host nests within the raiding range of their resident slavemaker nest, than to host nests outside the raiding range. This small‐scale population structure seems to be a result of polydomy (e.g. the occupation of several nests in close proximity by a single colony) and could have enabled the evolution of ‘rebellion’ by kin selection.     

Extreme life history plasticity and the evolution of invasive characteristics in a native ant

Disturbance resulting from urbanization is a leading cause of biotic homogenization worldwide. Native species are replaced with widespread non-native species and ants are among the world’s most notorious invaders. To date, all documented cases of ant invasions involve exotic introduced species that are spread around the world by human-mediated dispersal. I investigated the effect of urbanization on the evolution of invasive characteristics in a native ant species, the odorous house ant, Tapinoma sessile (Say). Colony social structure, life history traits, and the spatial pattern of nest distribution were compared by sampling T. sessile across a gradient of three distinct habitats: natural, semi-natural, and urban. Results demonstrate a remarkable transition in colony social and spatial structure and life history traits between natural and urban environments. In natural habitats, T. sessile colonies are comprised of small, monogyne (single queen), and monodomous (single nest) colonies. In urban areas, T. sessile often exhibit extreme polygyny and polydomy, form large supercolonies, and become a dominant pest. Results also suggest that urban T. sessile colonies may have a negative impact on native ant abundance and diversity. In the natural environment T. sessile coexisted with a wide array of other ant species, while very few ant species were present in the urban environment invaded by T. sessile. Habitat degradation and urbanization can lead to extreme changes in social and spatial colony structure and life history traits in a native ant species and can promote the evolution of invasive characteristics such as polygyny, polydomy, and supercolonial colony structure.     

Ecological consequences of colony structure in dynamic ant nest networks

Access to resources depends on an individual's position within the environment. This is particularly important to animals that invest heavily in nest construction, such as social insects. Many ant species have a polydomous nesting strategy: a single colony inhabits several spatially separated nests, often exchanging resources between the nests. Different nests in a polydomous colony potentially have differential access to resources, but the ecological consequences of this are unclear. In this study, we investigate how nest survival and budding in polydomous wood ant (Formica lugubris) colonies are affected by being part of a multi‐nest system. Using field data and novel analytical approaches combining survival models with dynamic network analysis, we show that the survival and budding of nests within a polydomous colony are affected by their position in the nest network structure. Specifically, we find that the flow of resources through a nest, which is based on its position within the wider nest network, determines a nest's likelihood of surviving and of founding new nests. Our results highlight how apparently disparate entities in a biological system can be integrated into a functional ecological unit. We also demonstrate how position within a dynamic network structure can have important ecological consequences.     

The population biology ofTapinoma minutum (Hymenoptera: Formicidae) in Australia

Summary Nests ofTapinoma minutum were collected and mapped from a wet sclerophyll forest in New South Wales during the spring, summer, and fall seasons. Queen number was variable, indicating the population is both facultatively polygynous and polydomous. Electrophoretic data from three polymorphic enzymes revealed that relatedness among workers conformed to the Hamiltonian expectation of 0.75. Colony boundaries were inferred from electrophoretic data synthesized with nest spatial locations. For this species colonies were composed of at most three nests; this simple pattern of polydomy suggests it has a recent origin in this population. The pattern of facultative polygyny may be linked to an apparent high rate of colony orphaning.     

Seasonal polydomy in a polygynous supercolony of the odorous house ant,Tapinoma sessile

Abstract 1. The odorous house ant, Tapinoma sessile, is a native ant species common throughout North America. In its natural habitat, T. sessile is a low‐key species that consists of small colonies. In invaded urban areas, T. sessile exhibits extreme polygyny and polydomy and becomes a dominant invasive pest. 2. The current study examined: (i) the density, persistence, and the spatial distribution of nests in a large supercolony of T. sessile, (ii) trail abundance and overall colony connectivity as facilitated by the network of trails, (iii) the abundance and the spatial distribution of competing ant species, and (iv) the effect of environmental factors on the number and distribution of T. sessile nests. 3. A distinct pattern of seasonal polydomy was observed, whereby the colony undergoes an annual fission‐fusion cycle. The colony occupies one or a few nests during the winter, experiences rapid exponential growth in the spring to colonize available nesting sites, reaches maximum nest density in the summer, and again coalesces in the winter, returning to the same winter location year after year. The trails show spatio‐temporal variation as well, depending on the location of nesting and foraging sites. Furthermore, nest movements may be driven by soil microclimate and proximity to man‐made structures. 4. In total, 119 ant nests were discovered in a 3.15 ha plot, 90 (76%) of which belonged to T. sessile. Tapinoma sessile exhibited strong colony connectivity as 78/90 (87%) of nests were connected to at least one other nest by a trail. Mean persistence time for T. sessile nests was 133 ± 5 days. 5. Results indicate that T. sessile is a highly adaptable native ant species that exhibits a high degree of flexibility in its colony social structure. A high degree of polygyny and polydomy may contribute to its ecological dominance and pest status in urban environments.     

Low intraspecific aggression level in the polydomous and facultative polygynous ant Ectatomma tuberculatum

Nestmate recognition is a key feature of social insects, as it preserves colony integrity. However, discrimination of non‐nestmates and nestmate recognition mechanisms are highly variable according to species and social systems. Here, we investigated the intraspecific level of aggression in the facultative polygynous and polydomous ant, Ectatomma tuberculatum Olivier (Hymenoptera: Formicidae: Ectatomminae), in a population with a strong genetic structure. We found that the intraspecific level of aggression was generally low in this population of E. tuberculatum. However, the level of aggression was significantly correlated with the geographical distance, suggesting that both genetic and environmental cues could be involved in nestmate recognition and discrimination mechanisms. Moreover, polydomy was confirmed by the absence of aggression between workers from nests at a distance of 3 m, while the level of aggression was significantly higher between workers from nests separated by a distance of 10 or 1300 m. Field experiments showed that the low level of aggression between neighbouring colonies was associated with shared foraging areas, which could suggest that familiarization processes may occur in this species. We propose that the particular social organization of this species, with secondary polygyny, polydomy, and budding, may have favoured a high acceptance threshold, because of the low probability of interactions with unrelated conspecifics, the high cost of erroneously rejecting nestmates, and the low cost of accepting non‐nestmate workers. The resulting open recognition system can thus allow privileged relationships between neighbouring colonies and promote the ecological dominance of E. tuberculatum in the mosaic of arboreal ants.     

The ecological benefits of larger colony size may promote polygyny in ants

How polygyny evolved in social insect societies is a long‐standing question. This phenomenon, which is functionally similar to communal breeding in vertebrates, occurs when several queens come together in the same nest to lay eggs that are raised by workers. As a consequence, polygyny drastically reduces genetic relatedness among nestmates. It has been suggested that the short‐term benefits procured by group living may outweigh the costs of sharing the same nesting site and thus contribute to organisms rearing unrelated individuals. However, tests of this hypothesis are still limited. To examine the evolutionary emergence of polygyny, we reviewed the literature to build a data set containing life‐history traits for 149 Palearctic ant species and combined this data set with a reconstructed phylogeny. We show that monogyny is the ancestral state and that polygyny has evolved secondarily and independently throughout the phylogenetic tree. The occurrence of polygyny is significantly correlated with larger colony size, dependent colony founding and ecological dominance. Although polydomy (when a colony simultaneously uses several connected nests) tends to occur more frequently in polygynous species, this trend is not significant when phylogenetic history is accounted for. Overall, our results indicate that polygyny may have evolved in ants in spite of the reduction in nestmate relatedness because large colony size provides immediate ecological advantages, such as the more efficient use of temporal food resources. We suggest that the competitive context of ant communities may have provided the conditions necessary for the evolution of polygyny in some clades.     

The Relationship between Canopy Cover and Colony Size of the Wood Ant Formica lugubris - Implications for the Thermal Effects on a Keystone Ant Species

Climate change may affect ecosystems and biodiversity through the impacts of rising temperature on species’ body size. In terms of physiology and genetics, the colony is the unit of selection for ants so colony size can be considered the body size of a colony. For polydomous ant species, a colony is spread across several nests. This study aims to clarify how climate change may influence an ecologically significant ant species group by investigating thermal effects on wood ant colony size. The strong link between canopy cover and the local temperatures of wood ant’s nesting location provides a feasible approach for our study. Our results showed that nests were larger in shadier areas where the thermal environment was colder and more stable compared to open areas. Colonies (sum of nests in a polydomous colony) also tended to be larger in shadier areas than in open areas. In addition to temperature, our results supported that food resource availability may be an additional factor mediating the relationship between canopy cover and nest size. The effects of canopy cover on total colony size may act at the nest level because of the positive relationship between total colony size and mean nest size, rather than at the colony level due to lack of link between canopy cover and number of nests per colony. Causal relationships between the environment and the life-history characteristics may suggest possible future impacts of climate change on these species.     

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.