The allocation of foragers in red wood ants

Abstract. 1. We studied how colonies of the red wood ant, Formica polyctena , adjust the numbers of foragers allocated to different foraging trails. In a series of field experiments, foragers were marked and transferred from one nest to another, related nest, where they joined the foraging force. Transferred workers acted as a reserve of uncommitted, available foragers.
2. Previous work shows that each individual forager habitually uses one trail. We found that for an uncommitted forager, the influence of recruitment initially is stronger than that of directional fidelity. Transferred workers were likely to use trails leading to new food sources. When transferred to a new nest, foragers were not likely to use a trail in the same direction as their original trail in the donor nest.
3. After a week, transferred foragers tended to develop route fidelity. Even after bait was no longer present, they continued to use the trail that had formerly led to a bait source.
4. We examined how colonies adjust numbers on a trail by experimentally depleting some trails. Colonies usually did not compensate for depletion: foragers were not recruited to depleted trails.
5. In general, the dynamics of foraging in this species facilitate a consistent foraging effort rather than rapid adjustments of forager allocation.     

An innate template for enemy recognition in red wood ants

The hypothesis of the innate template for perception and recognition of the enemy image in red wood ants Formica aquilonia Yarr. was tested by initiating conflicts between ants and predatory ground beetles, their competitors for space. Live beetles and their models with different characters were used. In nature, ants respond selectively to such features of competitors as dark coloration, the presence of “outgrowths” (legs, antennae), body symmetry, the rate of movement, and scent. Comparison of behavior of ants from natural colonies and “naive” (laboratory reared) ones showed that ants having no experience of encounters with competitors responded aggressively to an integral and sufficiently realistic enemy image. This suggests that red wood ants possess an innate template for recognition of potential competitors. At the same time, the ability to single out the key features and complete the integral image seems to require accumulation of experience.     

Spatial distribution of red wood ants and scientific basis of their protection

The results of long-term research on spatial distribution of red wood ants in the Urals and Western Siberia are presented. It was shown that large complexes counting tens and hundreds of nests clearly tended to form aggregations, or zones of high density. These zones are extended latitudinally and arranged in a striate pattern, with stripes about 15 km wide, about 100 km long, and positioned 100–120 km apart. The possibility of using these trends for the protection of red wood ants and for future research is discussed.     

Immigration and formation of mixed colonies in red wood ants (Hymenoptera,Formicidae)

Long-term field studies of the composition and spatial structure of settlements of ants of the Formica rufa group were carried out in two regions of Russia (Moscow and Arkhangelsk provinces). Fragmentation of damaged nests followed by reintegration of the fragments is the main way of formation of mixed colonies of ants from different nests (including different species). The principal factor of nest fragmentation is their damage by wild boars, bears, and in some localities, by poachers. The formation of mixed nests and nest complexes with participation of different Formica species was observed. They are formed by joining the ants from several damaged nests or by a colony from a destroyed nest immigrating into an intact one. Regular damage of many nests leads to the formation of broad zones of mixed colonies. The mixed colonies including 2–3 species of wood ants have recently become common. The phenomenon of mixed colonies raises a question as to the relative importance of two basic principles (sociality and specific identity) in the life of ant societies and demonstrates the priority of the social principle.     

Field tests of interspecific competition in ant assemblages: revisiting the dominant red wood ants

1. There has been considerable debate on the importance of competition in ecological communities, but its importance in structuring ant assemblages has often been uncritically accepted. Here, we briefly review field experiments examining competition in ant assemblages and use a removal experiment to test the effect of the classical territorial dominant ant, Formica aquilonia. Ants of this species group are thought to structure communities through a dominance hierarchy. 2. First, we used pitfall traps to compare the abundance of other ants in replicated sites with low and high densities of F. aquilonia. We found differences in community composition, in particular, Camponotus herculeanus was more common in low-density sites, in accordance with predictions. Differences in ant assemblages were not owing to differences in measured habitat variables. 3. We removed F. aquilonia from a set of high-density sites, using physical and chemical methods, and repeated these procedures at procedural control sites. One year after removal, abundances of F. aquilonia at removal sites were similar to those at low-density sites. However, the composition of other species did not change in response to F. aquilonia removal. Replication rates were identical in the mensurative and experimental components of this study, so this is unlikely to be owing to the analysis being insufficiently powerful. 4. We suggest three possibilities for the lack of difference. First, the study may have been too short term or small scale to detect differences. However, previous studies have shown effects on smaller spatial- and temporal-scales. Second, priority effects may be important in the successful colonisation by F. aquilonia. Thirdly, boreal ant assemblages may be too depauperate for redundancy in ecological roles and for competition to play an important structuring role. 5. We thus recommend that long-term large-scale experiments be considered essential if we are to distinguish between competing hypotheses in community ecology.     

Forestry alters foraging efficiency and crop contents of aphid-tending red wood ants, Formica aquilonia

Forest management alters species behaviours, distributions and interactions. To evaluate forestry effects on ant foraging performance, we compared the quality and quantity of honeydew harvested by ants among clear-cuts, middle-aged and mature spruce-dominated stands in boreal forests in Sweden. Honeydew quality was examined using honeydew collected by squeezing the gasters of laden Formica aquilonia workers. We used fifteen laden individuals at each study site (four replicates of each stand age) and analysed honeydew chemical composition with gas chromatography-mass spectroscopy. To compare the quantity of honeydew collected by individual ants, we collected and weighed five ants moving up and five ants moving down each of ten trees at the twelve sites (totally 1200 ants). The concentration of trehalose in honeydew was lower in clear-cuts compared with middle aged and mature stands, and similar trends were shown for sucrose, raffinose and melezitose, indicating poorer honeydew quality on clear cuts. Concentrations of the amino acid serine were higher on clear-cuts. The same trend occurred for glutamine, suggesting that increased N-uptake by the trees after clear cutting is reflected in the honeydew of aphids. Ants in mature stands had larger heads and carried proportionally more honeydew and may therefore be more efficient foragers. Human alternation of habitats through clear-cutting thus affects food quality and worker condition in F. aquilonia. This is the first study to show that honeydew quality is affected by anthropogenic disturbances, likely contributing to the reduction in size and abundance of F. aquilonia workers and mounds after clear cutting.     

An experimental study of behavioral mechanisms of relationships between red wood ants and ground beetles

Behavioral mechanisms regulating the competitive relations between the red wood ant Formica aquilonia and the ground beetles Carabus regalis, Pterostichus melanarius, P. magus, P. oblongopunctatus, Harpalus smaragdinus, and Amara nitida were studied by artificially initiated collisions between living insects as well as with the use of imitation models. Members of different functional groups within an ant family (aphid-milkers, hunters, and guards) behaved differently towards beetles. Active ants were shown to respond selectively to different features of the possible competitors, such as coloration, the presence of “appendages” (legs, antennae), body symmetry, rate of movement, and scent. Field and laboratory experiments demonstrated the ability of beetles to avoid collisions with active ants. The scent of anthill material attracted ground beetles, which consumed dead ants. The gained individual experience may allow the beetles to use supplementary forage resources in the territories controlled by ants. The flexible tactical patterns facilitate spatial segregation of ground beetles and ants in the same territory and result in a more complete utilization of food resources.     

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

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

Snapshot memories and landmark guidance in wood ants

Insects are thought to pinpoint a place by using memorized "snapshots," i.e., two-dimensional retinotopic views of the surrounding landmarks recorded when at the place (reviewed in ). Insects then reach the place by moving until their current view matches their snapshot. To determine when snapshots are recalled, and how differences between view and snapshot are translated into appropriate movements, we analyzed the approaches of wood ants to a feeding site that was located in the center of an array of two or three cylinders. In ants, contrary to flying hymenopterans, body orientation and direction of travel are collinear, so that an ant approaching an object always looks at it with frontal visual field. On their way to a food site, ants fixated and approached a cylinder predominantly when its angular size was smaller than when viewed from the food site. This finding implies that ants store snapshots at this place while fixating landmarks with frontal retina, so simplifying the later alignment of snapshots with their current view. It also means that ants recall snapshots well in advance of reaching the place. Although snapshots are centered on a landmark, we show that they extend at least 120 degrees into the periphery.     

Behavior of red wood ants (Hymenoptera,Formicidae) during interaction with different symbiont partners

A comparative analysis of the behavior of Formica polyctena Först during interaction with different symbionts (free-living aphids Aphis grossulariae Kalt. and hidden larvae of the sawfly Blasticotoma filiceti Klug) was carried out. Red wood ants demonstrate different levels of functional differentiation in relatively constant groups of foragers collecting honeydew. A deep “professional” specialization with clear division of a number of tasks among foragers was studied in groups of ants tending aphids. Four professional groups of foragers with different tasks were revealed: “shepherds,” “guards,” “transporters,” and “scouts” (or “coordinators”). The groups of foragers caring for sawfly larvae mainly consist of unspecialized ants. Only few ants (about 5%) remain on duty on the fern plant near B. filiceti larvae and protect the food resource from competitors, especially from other ants. In addition, the ants demonstrate simpler behavior while collecting the larval excretion, resembling that at the sugar feeders. On the whole, the behavior of red wood ants is rather flexible. The level of functional differentiation in groups of foragers collecting honeydew is determined not only by the colony size and requirements but by the nature of their interaction with trophobionts, particularly, by the possibility of direct contact.     

Within territory abundance of red wood ants Formica rufa is associated with the body condition of nestlings in the Eurasian treecreeper Certhia familiaris

Studies on individual reproductive success in relation to interspecific competition between distantly related taxa are scarce. We studied whether the abundance of red wood ants Formica rufa -group is related to the breeding habitat selection, fecundity and offspring quality in the Eurasian treecreeper Certhia familiaris, an old-growth forest passerine. The nest-box occupancy data were gathered over a five-year study period, whereas the breeding performance analyses were based on a two-year data set. The abundance of wood ants, measured within 50  m around the nest-boxes, was not related to nest-box occupancy rate, fecundity or the physiological stress of nestlings. In contrast, the abundance of wood ants was negatively related to the offspring quality and life-history traits, such as lowered body mass, subcutaneous fat reserves, and tarsus length at fledging. Our results suggest that exploitative competition between distantly related taxa may have considerable and adverse influences on nestling quality as measured by body mass, subcutaneous, fat and tarsus length. Red wood ants may decrease the fitness of treecreepers as the lower body condition of nestlings has the potential to impair recruitment into the breeding population and, additionally, impair the future reproductive effort.     

Scene perception and the visual control of travel direction in navigating wood ants

This review reflects a few of Mike Land''s many and varied contributions to visual science. In it, we show for wood ants, as Mike has done for a variety of animals, including readers of this piece, what can be learnt from a detailed analysis of an animal''s visually guided eye, head or body movements. In the case of wood ants, close examination of their body movements, as they follow visually guided routes, is starting to reveal how they perceive and respond to their visual world and negotiate a path within it. We describe first some of the mechanisms that underlie the visual control of their paths, emphasizing that vision is not the ant''s only sense. In the second part, we discuss how remembered local shape-dependent and global shape-independent features of a visual scene may interact in guiding the ant''s path.     

Visual cues for the retrieval of landmark memories by navigating wood ants

BACKGROUND: Even on short routes, ants can be guided by multiple visual memories. We investigate here the cues controlling memory retrieval as wood ants approach a one- or two-edged landmark to collect sucrose at a point along its base. In such tasks, ants store the desired retinal position of landmark edges at several points along their route. They guide subsequent trips by retrieving the appropriate memory and moving to bring the edges in the scene toward the stored positions. RESULTS: The apparent width of the landmark turns out to be a powerful cue for retrieving the desired retinal position of a landmark edge. Two other potential cues, the landmark's apparent height and the distance that the ant walks, have little effect on memory retrieval. A simple model encapsulates these conclusions and reproduces the ants' routes in several conditions. According to this model, the ant stores a look-up table. Each entry contains the apparent width of the landmark and the desired retinal position of vertical edges. The currently perceived width provides an index for retrieving the associated stored edge positions. The model accounts for the population behavior of ants and the idiosyncratic training routes of individual ants. DISCUSSION: Our results imply binding between the edge of a shape and its width and, further, imply that assessing the width of a shape does not depend on the presence of any particular local feature, such as a landmark edge. This property makes the ant's retrieval and guidance system relatively robust to edge occlusions.     

Ant assemblages in the taiga biome: testing the role of territorial wood ants

Summary Ants were collected with sets of pitfall traps in four coniferous-forest habitats in southern Finland. A three-level competition hierarchy concept was used to generate predictions on ant community structure. The levels of the hierarchy, and the respective predictions, from top to bottom were: (1) The dominant territorial wood ants (Formica rufa-group species), expected to exclude each other. (2) The other aggressive species, likely to be excluded by the F. rufa-group. (3) The submissive species, non-aggressive and defending only their nest, and thus likely to coexist with the dominants but in reduced numbers. As expected, the species of the F. rufa-group excluded each other, and the species number of the other aggressive ants was significantly cut down in the presence of the F. rufa-group. The aggressive species F. sanguinea and Camponotus herculeanus showed complementary occurrences with the F. rufa-group, and Lasius niger reduced occurrences. The number of the submissive species was not significantly affected by the F. rufa-group. However, pairwise correlation coefficients were significantly more often negative than positive between presence of the F. rufa-group and average proportion of pitfalls per set with a submissive species, each analyzed in turn. The result indicates that the F. rufa-group also reduced the colony densities of the submissive species. We conclude that in the taiga biome territorial wood ants are, after adjusting for physical vicissitudes of the environment, the major structuring force of ant species assemblages.     

Fight tactics in wood ants: individuals in smaller groups fight harder but die faster

When social animals engage in inter-group contests, the outcome is determined by group sizes and individual masses, which together determine group resource-holding potential ('group RHP'). Individuals that perceive themselves as being in a group with high RHP may receive a motivational increase and increase their aggression levels. Alternatively, individuals in lower RHP groups may increase their aggression levels in an attempt to overcome the RHP deficit. We investigate how 'group RHP' influences agonistic tactics in red wood ants Formica rufa. Larger groups had higher total agonistic indices, but per capita agonistic indices were highest in the smallest groups, indicating that individuals in smaller groups fought harder. Agonistic indices were influenced by relative mean mass, focal group size, opponent group size and opponent group agonistic index. Focal group attrition rates decreased as focal group relative agonistic indices increased and there was a strong negative influence of relative mean mass. The highest focal attrition rates were received when opponent groups were numerically large and composed of large individuals. Thus, fight tactics in F. rufa seem to vary with both aspects of group RHP, group size and the individual attributes of group members, indicating that information on these are available to fighting ants.     

Trail system in red wood ants (<Emphasis Type="Italic">Formica rufa</Emphasis> Group) under recreation press

The influence of recreation press on the forager trail system in red wood ants was examined. The model settlement of these ants was situated in green moss-pine forest, comprising areas with a different degree of the recreation press. In 2007, trail systems of 36 anthills were mapped (a total of 136 trails). In 2009, two nests from each group at each digression stage were selected as model objects. These nests possessed close to average values of the cupola diameter (d) and the number of trophic trails (n _c ). In these 6 model nests, trail systems (a total of 33 trails) were re-mapped and the strength of columns was estimated. It had been found that the recreation press decreases the population number in families of red wood ants. Trophic trails become shorter, their structure is simplified, and the number of trees with aphid colonies decreases. When the size of the family decreases owing to irregular decrease in the population number of separate columns, columns possessing non-ramified (less “efficient”) trails disappear first. Weakening of the family decreases the mobility of the trail network; trees with aphid colonies become more important.     

Terminating red imported fire ants using Cinnamomum osmophloeum leaf essential oil

Eleven compounds from indigenous cinnamon (Cinnamomum osmophloeum) leaf essential oil were identified by GC-MS and the dominant constituent was trans-cinnamaldehyde (79.85%). The toxicity of leaf essential oil and trans-cinnamaldehyde were then determined to study their effectiveness in controlling the red imported fire ant, Solenopsis invicta Buren. The results of the toxicity tests indicated that both the indigenous cinnamon leaf essential oil and trans-cinnamaldehyde had an excellent inhibitory effect in controlling the red imported fire ant. The LT(50) values for both 2% leaf essential oil and 2% trans-cinnamaldehyde after open exposure were 105.0min and 32.2min; after close exposure were 18.5min and 21.2min, respectively.     

Fire ant-detecting canines: a complementary method in detecting red imported fire ants

In this investigation, detection dogs are trained and used in identifying red imported fire ants, Solenopsis invicta Buren, and their nests. The methodology could assist in reducing the frequency and scope of chemical treatments for red imported fire ant management and thus reduce labor costs and chemical use as well as improve control and quarantine efficiency. Three dogs previously trained for customs quarantine were retrained to detect the scents of red imported fire ants. After passing tests involving different numbers of live red imported fire ants and three other ant species--Crematogaster rogenhoferi Mayr, Paratrechina longicornis Latreille, and Pheidole megacephala F.--placed in containers, ajoint field survey for red imported fire ant nests by detection dogs and bait traps was conducted to demonstrate their use as a supplement to conventional detection methods. The most significant findings in this report are (1) with 10 or more red imported fire ants in scent containers, the dogs had >98% chance in tracing the red imported fire ant. Upon the introduction of other ant species, the dogs still achieved on average, a 93% correct red imported fire ant indication rate. Moreover, the dogs demonstrated great competence in pinpointing emerging and smaller red imported fire ant nests in red imported fire ant-infested areas that had been previously confirmed by bait trap stations. (2) Along with the bait trap method, we also discovered that approximately 90% of red imported fire ants foraged within a distance of 14 m away from their nests. The results prove detection dogs to be most effective for red imported fire ant control in areas that have been previously treated with pesticides and therefore containing a low density of remaining red imported fire ant nests. Furthermore, as a complement to other red imported fire ant monitoring methods, this strategy will significantly increase the efficacy of red imported fire ant control in cases of individual mount treatment.