Trail using ants follow idiosyncratic routes in complex landscapes.

A large volume of research on individually navigating ants has shown how path integration and visually guided navigation form a major part of the ant navigation toolkit for many species and are sufficient mechanisms for successful navigation. One of the behavioural markers of the interaction of these mechanisms is that experienced foragers develop idiosyncratic routes that require that individual ants have personal and unique visual memories that they use to guide habitual routes between the nest and feeding sites. The majority of ants, however, inhabit complex cluttered environments and social pheromone trails are often part of the collective recruitment, organisation and navigation of these foragers. We do not know how individual navigation interacts with collective behaviour along shared trails in complex natural environments. We thus asked here if wood ants that forage through densely cluttered woodlands where they travel along shared trails repeatedly follow the same routes or if they choose a spread of paths within the shared trail. We recorded three long homing trajectories of 20 individual wood ants in their natural woodland habitat. We found that wood ants follow idiosyncratic routes when navigating along shared trails through highly complex visual landscapes. This shows that ants rely on individual memories for habitual route guidance even in cluttered environments when chemical trail information is available. We argue that visual cues are likely to be the dominant sensory modality for the idiosyncratic routes. These experiments shed new light on how ants, or insects in general, navigate through complex multimodal environments.

Semipermeable species boundaries create opportunities for gene flow and adaptive potential.

Hybridisation and gene flow can have both deleterious and adaptive consequences for natural populations and species. To better understand the extent of hybridisation in nature and the balance between its beneficial and deleterious outcomes in a changing environment, information on naturally hybridising nonmodel organisms is needed. This requires the characterisation of the structure and extent of natural hybrid zones. Here, we study natural populations of five keystone mound-building wood ant species in the Formica rufa group across Finland. No genomic studies across the species group exist, and the extent of hybridisation and genomic differentiation in sympatry is unknown. Combining genome-wide and morphological data, we demonstrate more extensive hybridisation than was previously detected between all five species in Finland. Specifically, we reveal a mosaic hybrid zone between Formica aquilonia, F. rufa and F. polyctena, comprising further generation hybrid populations. Despite this, we find that F. rufa, F. aquilonia, F. lugubris and F. pratensis form distinct gene pools in Finland. We also find that hybrids occupy warmer microhabitats than the nonadmixed populations of cold-adapted F. aquilonia, and suggest that warm winters and springs, in particular, may benefit hybrids over F. aquilonia, the most abundant F. rufa group species in Finland. In summary, our results indicate that extensive hybridisation may create adaptive potential that could promote wood ant persistence in a changing climate. Additionally, they highlight the potentially significant ecological and evolutionary consequences of extensive mosaic hybrid zones, within which independent hybrid populations face an array of ecological and intrinsic selection pressures.

Impact of central complex lesions on innate and learnt visual navigation in ants.

Wood ants are excellent navigators, using a combination of innate and learnt navigational strategies to travel between their nest and feeding sites. Visual navigation in ants has been studied extensively, however, we have little direct evidence for the underlying neural mechanisms. Here, we perform lateralized mechanical lesions in the central complex (CX) of wood ants, a midline structure known to allow an insect to keep track of the direction of sensory cues relative to its own orientation and to control movement. We lesioned two groups of ants and observed their behaviour in an arena with a large visual landmark present. The first group of ants were naïve and when intact such ants show a clear innate attraction to the conspicuous landmark. The second group of ants were trained to aim to a food location to the side of the landmark. The general heading of naïve ants towards a visual cue was not altered by the lesions, but the heading of ants trained to a landmark adjacent food position was affected. Thus, CX lesions had a specific impact on learnt visual guidance. We also observed that lateralised lesions altered the fine details of turning with lesioned ants spending less time turning to the side ipsilateral of the lesion. The results confirm the role of the CX in turn control and highlight its important role in the implementation of learnt behaviours that rely on information from other brain regions.

Whole-genome analysis of multiple wood ant population pairs supports similar speciation histories, but different degrees of gene flow, across their European ranges.

The application of demographic history modelling and inference to the study of divergence between species has become a cornerstone of speciation genomics. Speciation histories are usually reconstructed by analysing single populations from each species, assuming that the inferred population history represents the actual speciation history. However, this assumption may not be met when species diverge with gene flow, for example, when secondary contact may be confined to specific geographic regions. Here, we tested whether divergence histories inferred from heterospecific populations may vary depending on their geographic locations, using the two wood ant species Formica polyctena and F. aquilonia. We performed whole-genome resequencing of 20 individuals sampled in multiple locations across the European ranges of both species. Then, we reconstructed the histories of distinct heterospecific population pairs using a coalescent-based approach. Our analyses always supported a scenario of divergence with gene flow, suggesting that divergence started in the Pleistocene (c. 500 kya) and occurred with continuous asymmetrical gene flow from F. aquilonia to F. polyctena until a recent time, when migration became negligible (2-19 kya). However, we found support for contemporary gene flow in a sympatric pair from Finland, where the species hybridise, but no signature of recent bidirectional gene flow elsewhere. Overall, our results suggest that divergence histories reconstructed from a few individuals may be applicable at the species level. Nonetheless, the geographical context of populations chosen to represent their species should be taken into account, as it may affect estimates of migration rates between species when gene flow is spatially heterogeneous.

Challenges and a call to action for protecting European red wood ants.

Red wood ants (RWAs) are a group of keystone species widespread in temperate and boreal forests of the Northern Hemisphere. Despite this, there is increasing evidence of local declines and extinctions. We reviewed the current protection status of RWAs throughout Europe and their International Union for the Conservation of Nature (IUCN) threat classification. Only some RWA species have been assessed at a global scale, and not all national red lists of the countries where RWAs are present include these species. Different assessment criteria, inventory approaches, and risk categories are used in different countries, and data deficiency is frequent. Legislative protection is even more complex, with some countries protecting RWAs implicitly together with the wildlife fauna and others explicitly protecting the whole group or particular species. This complexity often occurs within countries, for example, in Italy, where, outside of the Alps, only the introduced species are protected, whereas the native species, which are in decline, are not. Therefore, an international, coordinated framework is needed for the protection of RWAs. This first requires that the conservation target should be defined. Due to the similar morphology, complex taxonomy, and frequent hybridization, protecting the entire RWA group seems a more efficient strategy than protecting single species, although with a distinction between autochthonous and introduced species. Second, an update of the current distribution of RWA species is needed throughout Europe. Third, a protection law cannot be effective without the collaboration of forest managers, whose activity influences RWA habitat. Finally, RWA mounds offer a peculiar microhabitat, hosting a multitude of taxa, some of which are obligate myrmecophilous species on the IUCN Red List. Therefore, RWAs' role as umbrella species could facilitate their protection if they are considered not only as target species but also as providers of species-rich microhabitats.

Las hormigas rojas de la madera (HRM) conforman un grupo de especies clave con amplia distribución en los bosques templados y boreales del Hemisferio Norte. A pesar de lo anterior, cada vez hay más evidencia de su declinación y extinción local. Revisamos el estado actual de protección de las HRM en toda Europa y su clasificación en la Lista Roja de la Unión Internacional para la Conservación de la Naturaleza (UICN). Sólo se han evaluado algunas especies de HRM a escala mundial y no todas las listas rojas nacionales de los países con presencia de HRM incluyen a estas especies. Los diferentes países usan criterios de evaluación, estrategias de inventario y categorías de riesgo distintos, además de que la información deficiente es habitual. La protección legislativa es todavía más compleja pues algunos países protegen implícitamente a las HRM junto con la fauna silvestre y otros protegen explícitamente a todo el grupo o a una especie particular. Esta complejidad ocurre a menudo en los países (por ejemplo: Italia) en donde, fuera de los Alpes, sólo se protege a las especies introducidas, mientras a las especies nativas, que están declinando, no se les protege. Por lo tanto, se requiere un marco de trabajo internacional y coordinado para proteger a las HRM. Esto necesita primero que se defina el objetivo de conservación. Ya que las HRM tienen similitudes morfológicas, una taxonomía compleja e hibridación frecuente, la protección del grupo completo, con la distinción entre las especies autóctonas y las introducidas, parece ser una estrategia más eficiente que la protección de una sola especie. Segundo, se debe actualizar la distribución actual de las HRM en Europa. Tercero, una ley de protección no puede ser efectiva sin la colaboración de los gestores forestales, cuya actividad influye sobre el hábitat de las HRM Finalmente, los montículos de las HRM ofrecen un microhábitat peculiar pues hospedan a una multitud de taxones, algunos de los cuales son especies mirmecófilas obligadas presentes en la Lista Roja de la UICN. Así, el papel de las HRM como especie paraguas podría facilitar su protección si se les considera no sólo como especies diana sino también como proveedoras de microhábitats con riqueza de especies.

Innate visual attraction in wood ants is a hardwired behavior seen across different motivational and ecological contexts.

Ants are expert navigators combining innate and learnt navigational strategies. Whereas we know that the ants' feeding state segregates visual-navigational memories in ants navigating along a learnt route, it is an open question if the motivational state also affects the ants' innate visual preferences. Wood ant foragers show an innate attraction to conspicuous visual cues. These foragers inhabit cluttered woodland habitat and feed on honeydew from aphids on trees. Hence, the attraction to 'tree-like' objects might be an ecologically relevant behavior that is tailored to the wood ants' foraging ecology. Foragers from other ant species with different foraging ecologies show very different innate attractions. We investigated here the innate visual response of wood ant foragers with different motivational states, i.e., unfed or fed, as well as males that show no foraging activity. Our results show that ants from all three groups orient toward a prominent visual cue, i.e., this intrinsic visuomotor response is not context-dependent, but a hardwired behavior seen across different motivational and ecological contexts. Supplementary Information: The online version contains supplementary material available at 10.1007/s00040-022-00867-3.

Ant colony nest networks adapt to resource disruption.

Animal social structure is shaped by environmental conditions, such as food availability. This is important as conditions are likely to change in the future and changes to social structure can have cascading ecological effects. Wood ants are a useful taxon for the study of the relationship between social structure and environmental conditions, as some populations form large nest networks and they are ecologically dominant in many northern hemisphere woodlands. Nest networks are formed when a colony inhabits more than one nest, known as polydomy. Polydomous colonies are composed of distinct sub-colonies that inhabit spatially distinct nests and that share resources with each other. In this study, we performed a controlled experiment on 10 polydomous wood ant (Formica lugubris) colonies to test how changing the resource environment affects the social structure of a polydomous colony. We took network maps of all colonies for 5 years before the experiment to assess how the networks changes under natural conditions. After this period, we prevented ants from accessing an important food source for a year in five colonies and left the other five colonies undisturbed. We found that preventing access to an important food source causes polydomous wood ant colony networks to fragment into smaller components and begin foraging on previously unused food sources. These changes were not associated with a reduction in the growth of populations inhabiting individual nests (sub-colonies), foundation of new nests or survival, when compared with control colonies. Colony splitting likely occurred as the availability of food in each nest changed causing sub-colonies to change their inter-nest connections. Consequently, our results demonstrate that polydomous colonies can adjust to environmental changes by altering their social network.

Emergence of a complex movement pattern in an unfamiliar food place by foraging ants.

Although visual cues are essential for navigation in ants, few studies address movement dynamics in ants when they search and forage after finding food in an unfamiliar environment. Here I introduced Japanese wood ants to an unfamiliar food location by capturing individuals leaving their nest. The food was located at the centre of a straight, narrow, open-top channel. Next, I determined the segment lengths of the foraging paths of the ants between consecutive U-turns. I found that individuals travelled along characteristic and complex paths if they detected a visual landmark. This movement property was not detectable when individuals foraged in the channel without any visual landmarks. These results reveal the movement dynamics of ants when they encounter food in a novel place.

Widespread hybridization within mound-building wood ants in Southern Finland results in cytonuclear mismatches and potential for sex-specific hybrid breakdown.

Hybridization and gene flow between diverging lineages are increasingly recognized as common evolutionary processes, and their consequences can vary from hybrid breakdown to adaptive introgression. We have previously found a population of wood ant hybrids between Formica aquilonia and F. polyctena that shows antagonistic effects of hybridization: females with introgressed alleles show hybrid vigour, whereas males with the same alleles show hybrid breakdown. Here, we investigate whether hybridization is a general phenomenon in this species pair and analyse 647 worker samples from 16 localities in Finland using microsatellite markers and a 1200-bp mitochondrial sequence. Our results show that 27 sampled nests contained parental-like gene pools (six putative F. polyctena and 21 putative F. aquilonia) and all remaining nests (69), from nine localities, contained hybrids of varying degrees. Patterns of genetic variation suggest these hybrids arise from several hybridization events or, instead, have backcrossed to the parental gene pools to varying extents. In contrast to expectations, the mitochondrial haplotypes of the parental species were not randomly distributed among the hybrids. Instead, nests that were closer to parental-like F. aquilonia for nuclear markers preferentially had F. polyctena's mitochondria and vice versa. This systematic pattern suggests there may be underlying selection favouring cytonuclear mismatch and hybridization. We also found a new hybrid locality with strong genetic differences between the sexes similar to those predicted under antagonistic selection on male and female hybrids. Further studies are needed to determine the selective forces that act on male and female genomes in these newly discovered hybrids.

Distribution and physical traits of red wood ant mounds in a managed Rhodope mountains forest.

Red wood ants (RWA) are of great ecological importance for the forest ecosystem. Forestry practices, like clear-cutting, and trampling load, due to tourism, logging, and grazing stock, can greatly affect their colonies, disturbing their microhabitat. RWA in Greek forests have not been investigated so far. We herein report on the distribution and morphological traits of Formica lugubris mounds studied in Elatia forest (Rhodope mountains, Northern Greece), an all-aged managed mixed forest where selective logging practices are performed. Nearby vegetation, slope, canopy cover, shrub density, and distance from the nearest neighboring trees were also recorded. Mound density was shown to be much higher in this Greek forest compared to RWA mounds in other European-managed forests. Furthermore, we recorded a continuous nest establishment, despite forest management disturbances and trampling load. Our study suggests that single-tree selective forestry practices are essential for creating ideal microhabitats for the RWA and, therefore, for maintaining RWA populations.

Intraspecific Relationships and Nest Mound Shape Are Affected by Habitat Features in Introduced Populations of the Red Wood Ant Formica paralugubris.

Ants belonging to the Formica rufa group build large nest mounds, which aid their survival during severe winters. We investigated whether different environmental features of the habitats affected the nest mound shape and the population structure. We assessed the shape of all the nest mounds and mapped inter-nest trails connecting mounds for three imported populations of Formica paralugubris in three forest habitats: fir-dominated, beech-dominated, and a mixture of fir and beech. Single-nest mounds were averagely smaller and flatter in the beech-dominated forest, probably because of lighter building materials. Nonetheless, by summing the volumes of all interconnected nests, the size was similar among all three sites. In fir- and beech-dominated forests, large nests were also central in the networks, suggesting a central place foraging model with these nests as reference. We finally performed aggression tests, and found that aggressiveness was significantly higher among nests belonging to the same population than between populations. The results highlight the plasticity of the species to adapt nest and colony structure to different environments. Additionally, it appears that none of these populations is unicolonial, as observed in various alpine sites, there and the observed patterns of aggression are coherent with the 'nasty neighbor' effect.

Assessing Molecular Diversity in Native and Introduced Populations of Red Wood Ant Formica paralugubris.

The Formica rufa group comprises several ant species which are collectively referred to as "red wood ants" and play key roles in boreal forest ecosystems, where they are ecologically dominant and greatly influence habitat dynamics. Owing to their intense predatory activity, some of these species are used as biocontrol agents against several forest insect pests and for this aim in Italy, nearly 6000 ant nests were introduced from their native areas in the Alps to several Appeninic sites during the last century. In this work, we assessed and compared the genetic variability and structure of native and introduced populations of F. paralugubris, thus evaluating the extent of genetic drift that may have occurred since the time of introduction, using amplified fragment length polymorphism (AFLP) markers. PCR amplification with a fam_EcoRI-TAC/MseI-ATG primers combination produced a total of 147 scorable bands, with 17 identified as outlier loci. The genetic variation was higher in the introduced population compared to the native ones that, on the other hand, showed a higher diversity between nests. AMOVA results clearly pointed out that the overall genetic structure was dominated by among-worker variation, considering all populations, the Alpine vs. Apennine groups and the comparison among native and related introduced populations (all ranging between 77.84% and 79.84%). Genetic analyses unveiled the existence of six main different groups that do not entirely mirror their geographic subdivision, pointing towards a wide admixture between populations, but, at the same time, rapid diversification of some Apennine populations. Future studies based on high-throughput genomic methods are needed to obtain a thorough understanding of the effects of environmental pressure on the genetic structure and mating system of these populations.

Comparative Analysis of Facial Coloration between Introduced and Source Populations of the Red Wood Ant Formica paralugubris.

The variation in the typical black-reddish color of red wood ants (Formica rufa group) has been recently suggested as a good indicator of habitat quality, being dependent on environmental conditions. However, the relative contribution of external factors and heritability in shaping this trait is poorly investigated. In this study, we compared the facial coloration of workers from four introduced populations of Formica paralugubris with those of the two Alpine populations from which they had been taken. We used a Relative Warp Analysis to describe the variations in the shape of this trait. We expected each introduced population to be more similar to its population of origin if the color pattern was predominantly genetically determined. On the contrary, due to the considerable differences in habitat type and climate between the Alps and the Apennines, we expected to observe differences between the introduced population and their origin population if the coloration was mostly environmentally determined. With one exception that we discuss, the results showed that ants from the two source populations had different phenotypes, and that the introduced populations had a shape similar to the population of origin, suggesting a stable genetic background. Surprisingly, the habitat type seems to have a less clear effect, even if within-population differences suggest the influence of very localized environmental factors. Finally, we found that the facial coloration shape is affected by the ant's size, a result in line with previous studies.

Chance or Necessity-The Fungi Co-Occurring with Formica polyctena Ants.

Studies on carton nesting ants and domatia-dwelling ants have shown that ant-fungi interactions may be much more common and widespread than previously thought. Until now, studies focused predominantly on parasitic and mutualistic fungi-ant interactions occurring mostly in the tropics, neglecting less-obvious interactions involving the fungi common in ants' surroundings in temperate climates. In our study, we characterized the mycobiota of the surroundings of Formica polyctena ants by identifying nearly 600 fungal colonies that were isolated externally from the bodies of F. polyctena workers. The ants were collected from mounds found in northern and central Poland. Isolated fungi were assigned to 20 genera via molecular identification (ITS rDNA barcoding). Among these, Penicillium strains were the most frequent, belonging to eight different taxonomic sections. Other common and widespread members of Eurotiales, such as Aspergillus spp., were isolated very rarely. In our study, we managed to characterize the genera of fungi commonly present on F. polyctena workers. Our results suggest that Penicillium, Trichoderma, Mucor, Schwanniomyces and Entomortierella are commonly present in F. polyctena surroundings. Additionally, the high diversity and high frequency of Penicillium colonies isolated from ants in this study suggest that representatives of this genus may be adapted to survive in ant nests environment better than the other fungal groups, or that they are preferentially sustained by the insects in nests.

Mushroom Bodies Are Required for Learned Visual Navigation, but Not for Innate Visual Behavior, in Ants.

Visual navigation in ants has long been a focus of experimental study [1-3], but only recently have explicit hypotheses about the underlying neural circuitry been proposed [4]. Indirect evidence suggests the mushroom bodies (MBs) may be the substrate for visual memory in navigation tasks [5-7], while computational modeling shows that MB neural architecture could support this function [8, 9]. There is, however, no direct evidence that ants require MBs for visual navigation. Here we show that lesions of MB calyces impair ants' visual navigation to a remembered food location yet leave their innate responses to visual cues unaffected. Wood ants are innately attracted to large visual cues, but we trained them to locate a food source at a specific angle away from such a cue. Subsequent lesioning of the MB calyces using procaine hydrochloride injection caused ants to revert toward their innate cue attraction. Handling and saline injection control ants still approached the feeder. Path straightness of lesioned and control ants did not differ from each other but was lower than during training. Reversion toward the cue direction occurred irrespective of whether the visual cue was ipsi- or contralateral to the lesion site, showing this is not due simply to an induced motor bias. Monocular occlusion did not diminish ants' ability to locate the feeder, suggesting that MB lesions are not merely interrupting visual input to the calyx. The demonstrated dissociation between innate and learned visual responses provides direct evidence for a specific role of the MB in navigational memory.

First In Situ Identification of Ultradian and Infradian Rhythms, and Nocturnal Locomotion Activities of Four Colonies of Red Wood Ants ( Formica rufa-Group).

In situ activity patterns of 2 Formica rufa-group species ( F. pratensis; F. polyctena) were continuously studied at 4 different red wood-ant nests for 6 months in each of the years 2010, 2011, 2012, and 2016 and related to weather factors and variations in the Earth's magnetic field. The in situ activity patterns of both species were similarly periodic and exhibited ultradian, and short- and long infradian rhythms under natural LD conditions. Crepuscular and nocturnal activities shorter than or equal to 4 h were observed in both species, especially at the new moon and first quarter after the astronomical twilight in a period of darkness in fall. We hypothesize that local variability in the Earth's magnetic field affects these long-term activity patterns, whereas humidity and temperature were more strongly associated with ultradian rhythms (less than 20 h).

Can a Red Wood-Ant Nest Be Associated with Fault-Related CH4 Micro-Seepage? A Case Study from Continuous Short-Term In-Situ Sampling.

We measured methane (CH4) and stable carbon isotope of methane (δ13C-CH4) concentrations in ambient air and within a red wood-ant (RWA; Formica polyctena) nest in the Neuwied Basin (Germany) using high-resolution in-situ sampling to detect microbial, thermogenic, and abiotic fault-related micro-seepage of CH4. Methane degassing from RWA nests was not synchronized with earth tides, nor was it influenced by micro-earthquake degassing or concomitantly measured RWA activity. Two δ13C-CH4 signatures were identified in nest gas: -69‰ and -37‰. The lower peak was attributed to microbial decomposition of organic matter within the RWA nest, in line with previous observations that RWA nests are hot-spots of microbial CH4. The higher peak has not been reported in previous studies. We attribute this peak to fault-related CH4 emissions moving via fault networks into the RWA nest, which could originate either from thermogenic or abiotic CH4 formation. Sources of these micro-seepages could be Devonian schists, iron-bearing "Klerf Schichten", or overlapping micro-seepage of magmatic CH4 from the Eifel plume. Given the abundance of RWA nests on the landscape, their role as sources of microbial CH4 and biological indicators for abiotically-derived CH4 should be included in estimation of methane emissions that are contributing to climatic change.

Comparative larval ultramorphology of some myrmecophilous Aleocharinae (Coleoptera, Staphylinidae), with a first description of the larvae of Amidobiatalpa (Heer O, 1841) and Oxypodahaemorrhoa (Mannerheim C.G., 1830), associated with the Formicarufa species group.

The paper describes the external structures of the late larval stages of two Palearctic myrmecophilous staphylinids: Amidobiatalpa and Oxypodahaemorrhoa associated with the Formicarufa species group. This is the first-ever description of the larva of Amidobia, and the only complete, detailed account of the morphology of this developmental stage in the genus Oxypoda currently available. For the first time in these two genera, 13 and 10 larval diagnostic features, respectively, are proposed. Morphological differences have been established between known and the newly described larvae of five species (genera) of myrmecophilous and one non-myrmecophilous Aleocharinae, belonging to three tribes. Amidobiatalpa and O.haemorrhoa are probably typical, tiny predators, like most other Aleocharinae, including non-myrmecophilous ones. Being very small and highly mobile, they are ignored by worker ants. Not surprisingly, no particular larval morphological modifications were found to enable them to survive among ants. Such features have, however, evolved in the larvae of larger aleocharines, that is, those that are perceived by ants and are wholly integrated with their hosts in the ant nest (e.g. Lomechusa). This comparative analysis of the functional morphology of the larvae of known myrmecophilous Aleocharinae is a springboard to further such studies of these interesting insects.

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.

Internest food sharing within wood ant colonies: resource redistribution behavior in a complex system.

Resource sharing is an important cooperative behavior in many animals. Sharing resources is particularly important in social insect societies, as division of labor often results in most individuals including, importantly, the reproductives, relying on other members of the colony to provide resources. Sharing resources between individuals is therefore fundamental to the success of social insects. Resource sharing is complicated if a colony inhabits several spatially separated nests, a nesting strategy common in many ant species. Resources must be shared not only between individuals in a single nest but also between nests. We investigated the behaviors facilitating resource redistribution between nests in a dispersed-nesting population of wood ant Formica lugubris. We marked ants, in the field, as they transported resources along the trails between nests of a colony, to investigate how the behavior of individual workers relates to colony-level resource exchange. We found that workers from a particular nest "forage" to other nests in the colony, treating them as food sources. Workers treating other nests as food sources means that simple, pre-existing foraging behaviors are used to move resources through a distributed system. It may be that this simple behavioral mechanism facilitates the evolution of this complex life-history strategy.