Worker reproduction in the ant Formica fusca

We studied the kin conflict over male parentage in the ant Formica fusca. The conflict arises because each worker and queen is most related to her own sons and is thus predicted to lay eggs. Microsatellite analysis of eggs revealed that workers laid eggs in more than half the queenright experimental nests. Nevertheless, almost exclusively diploid offspring were reared in the presence of a queen. This also occurred when worker-laid haploid male eggs were experimentally introduced in to the nests. Because our experimental setup allowed us to exclude the possibility of queen policing, we conclude that worker laid eggs are removed by other workers, either as a response to their parentage or gender. Our results suggest that worker reproduction in F. fusca is ultimately an interplay of conflicts over male parentage and sex allocation and that both worker and self policing have roles as proximate mechanisms of resolution.     

Effects of nest size and dispersion on brood production in a North American population of wood ant Formica fusca (Hymenoptera: Formicidae)

We examined several key parameters of the population ecology of a North American population of Formica fusca (L.), including nest dispersion, colony size and brood production. Physical nest size was significantly correlated with colony size, and colony size, in turn, was significantly correlated with brood production. Sex allocation was male biased, although larger nests were more likely to produce reproductive female brood (gynes). Neither nest temperature nor moisture level was significantly correlated with brood production. Formica fusca nests in this population had a comparatively low average nearest-neighbor distance with a significantly even pattern of dispersion, which suggests relatively high intraspecific competition. However, nearest-neighbor distance was not significantly associated with either colony size or relative brood production.     

Interaction between the non-myrmecochorous herb Galium aparine and the ant Formica polyctena

Non-myrmecochorous plant species are able to colonize Formica polyctena mounds. Galium aparine has been chosen as a model to study this kind of ant-plant association. In order to understand benefits to ants and plants, we have studied removal rates of fruits by ant workers, and compared soil seed material, seedling pattern and development of plants of the nest versus forest.Ant workers remove fruits at very low rates; removal time for half of the sample of 10 seeds was on average 5 h. Removed fruits were usually lost within 20–30 cm distance of the ant route. The soil samples from the nests contained a higher number of fruits, than those from randomly chosen sites in the forest. Seeds start to germinate in late August; on the F. polyctena mounds significantly earlier than at surrounding places. The seedling density was higher within experimental quadrats located on mound sites especially in late September–early October. The highest density of seedlings was on the slope of the mound. Plants from mounds are significantly longer, have a higher number of nodes during all stages of development and a higher mortality rate than that at surrounding places.The interactions between G. aparine and F. polyctena ants are asymmetrical. In spite of higher mortality rate of seedlings on mounds, ants provide some benefits to the plant. Firstly, workers are additional dispersal agents for G. aparine fruits. Furthermore, the soil peculiarities of ant mounds are beneficial for seed germination and seedling growth. G. aparine does not provide ants with energetically valuable food like plants with elaiosome-bearing seeds. Moreover, the high density of seedlings and plants on the mounds shades the nest.     

Population studies on hill building wood-ants of the Formica rufa-group

Summary Population studies on hill building wood-ants of the Formica rufa-group.Total inventories of inhabited ant hills of the Formica rufa-group were carried out in a part of the Black Forest amounting to 1,640 ha (4,100 acres) near Freiburg im Breisgau on five occasions during the period 1966–1978. Nest densities of 12.7 to 19.1 per 100 ha were recorded. Mean annual nest mortality for the three most abundant species (F. rufa, F. polyctena, F. pratensis) was 21–33% (Table 2); it remained almost the same during all intervals between inventories for all age classes studied (Fig. 2). The mean annual nest natality varied between 31 and 51%. It was therefore concluded that changes in the numbers of ant hills (Fig. 1) resulted from nest natality rather than from the reasonably stable nest mortality. Nest size was assessed by height and diameter. The development of nest size showed a marked increase with age for F. rufa and F. polyctena reaching a maximum at 2–5 years. Mean life expectancy of small nests was markedly lower than that of larger nests (Fig. 3). Consequently in the large size classes the numbers of old nests were significantly greater than the numbers of newly founded nests (Fig. 4). Results are compared with those recorded in the literature for palearctic and nearctic species belonging to the Formica rufa-group.Supported by Ministerium für Ernährung, Landwirtschaft und Umwelt Baden-Württemberg (MELU) and Gesellschaft zur Förderung der Forst- und Holzwirtschaftlichen Forschung Freiburg     

Effects of reproductive timing and colony size on the survival, offspring colony size and drone production in the honey bee (Apis mellifera)

ABSTRACT. 1. The effects of colony size and time of reproduction on the survival and size of offspring colonies and on drone production were examined for honey bees, Apis mellifera L. Drone and worker production and survival of parental and offspring colonies were monitored following swarming. Also, the temporal patterns of drone emergence and availability of unmated queens were examined.
2. Colony size at swarming was positively correlated with the number of workers invested in offspring colonies and the number of queens produced. However, colony size at swarming was not correlated with the number of offspring colonies produced.
3. Swarm size was positively correlated with drone and worker production after swarms were hived. Worker production of hived swarms was positively correlated with colony survival. Offspring queens which inherited a parental nest survived longer than queens in either primary swarms or afterswarms, presumably due to the advantage of inheriting a nest.
4. Drone emergence peaked just prior to swarming, the time when unmated queens were available. High drone production by colonies initiated by swarms probably reflected an attempt to reproduce prior to winter. The probabilities of a second swarming cycle within the same year and of surviving the winter were low for colonies initiated from swarms.     

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

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

The size–distance relationship in the wood ant Formica rufa

1. The size–distance relationship among honeydew‐collecting foragers of the red wood ant Formica rufa was investigated. Within the colony territory, the size (as measured by head width) and fresh weight of samples of foragers were determined for ants ascending and descending trees near, and farther from, the central nest mound. 2. The mean size of the ants was significantly higher at far trees than at near trees in six out of the seven colonies investigated, confirming the general presence of the size–distance relationship. 3. In three colonies, a load–distance relationship was also found. For a given head width, honeydew‐carrying ants descending far trees were significantly heavier than those descending near trees (i.e. they were carrying heavier loads from trees farther away from the central nest mound). 4. This is the first time that both load–distance and size–distance relationships have been reported in foraging workers from the same ant colony. 5. The combined effects of these characteristics suggest that colony foraging efficiency is enhanced by far trees being visited by the larger workers that then return with heavier loads of honeydew.     

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.     

Responses of Two Potential Host Species (Formica gnava and Formica occulta) to Pupae of the Obligatory Slave-Making Ant, Polyergus breviceps (Hymenoptera: Formicidae)

Queens of the slave-making ant, Polyergus breviceps, take over nests of adult Formica workers when establishing new colonies. Although naïve to slave-maker brood, the usurped Formica rear Polyergus offspring and nests containing both host and parasite species forms. Host worker acceptance of parasite brood has been attributed to the similarity of brood tending signals between these closely related taxa and/or the presence of an attractive pheromone in the slave-maker brood. By presenting single-species groups of Formica occulta and Formica gnava (two host species of P. breviceps) with a choice of Formica pupae of both species or with a choice of P. breviceps pupae from both types of mixed-species nests, it seems that neither close phylogenetic relatedness nor an attractive brood odor alone can account for the propensity of host workers to adopt slave-maker pupae. Significantly greater numbers of P. breviceps pupae were adopted by enslaved workers than by free-living workers, and within the enslaved groups and the free-living F. gnava group, greater numbers of P. breviceps pupae were adopted if they were from nests where the host species was conspecific to workers used in tests. When presented with F. gnava and F. occulta pupae, Formica workers adopted conspecific pupae almost exclusively and ignored or consumed pupae of the other host species. Taken together, these results imply that P. breviceps pupae have both a species-specific odor and a general brood-tending pheromone, upon which a host odor may be imposed. The disparate requirements of immatures at different stages of development for cue specificity or generality in maintaining nest exclusivity and maximizing inclusive fitness are discussed.     

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

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

Multiple breeders, breeder shifts and inclusive fitness returns in an ant

In social insects, colonies may contain multiple reproductively active queens. This leads to potential conflicts over the apportionment of brood maternity, especially with respect to the production of reproductive offspring. We investigated reproductive partitioning in offspring females (gynes) and workers in the ant Formica fusca, and combined this information with data on the genetic returns gained by workers. Our results provide the first evidence that differential reproductive partitioning among breeders can enhance the inclusive fitness returns for sterile individuals that tend non-descendant offspring. Two aspects of reproductive partitioning contribute to this outcome. First, significantly fewer mother queens contribute to gyne (new reproductive females) than to worker brood, such that relatedness increases from worker to gyne brood. Second, and more importantly, adult workers were significantly more related to the reproductive brood raised by the colony, than to the contemporary worker brood. Thus, the observed breeder shift leads to genetic benefits for the adult workers that tend the brood. Our results also have repercussions for genetic population analyses. Given the observed pattern of reproductive partitioning, estimates of effective population size based on worker and gyne samples are not interchangeable.     

Wood ants are wood ants: deforestation causes population declines in the polydomous wood ant Formica aquilonia

Abstract.  1. One of the main themes in ecology is adaptation for survival in different habitats and the potential of the environment to regulate populations.
2. The effects of clear-cutting on nest-abandonment rate and local population sizes in the polydomous wood ant Formica aquilonia was studied, using uncut forest stands as controls.
3. The nest-abandonment rate was clearly higher in clear-cuts than in forest interiors. In clear-cuts, 39% of pre-deforestation nests and 73% of new bud-nests were abandoned 4–5 years after deforestation, whereas in forest interiors fewer than 2% of nests were abandoned at the same time period. Local population size decreased 30% in clear-cuts, but fewer than 2% in forest interiors.
4. The results demonstrate that despite modern logging practices in which mechanical harming of nest mounds is reduced, nest mounds are abandoned at high rate, and despite frequent establishment of new bud-nests, populations start to decline.
5. The likely reason for the high nest-abandonment rate in clear-cuts is a combination of changed abiotic conditions, resource limitation, and disturbed reproduction.
6. Species that are sensitive to changes in the size of habitat patch, such as F. aquilonia , likely are harmed by logging, even employing biodiversity oriented management practices. Hence there is a need for conservation actions that are based on the size of protection areas.     

Reproductive strategy of the slave antFormica podzolica relative to raiding efficiency of enslaver species

Summary Formica podzolica serves as host to slave-making ants in North America. We propose thatF. podzolica may respond to slavery by two alternative colony-growth and reproductive strategies depending on the raiding ability of the slavemaker: (1) Rapid colony growth at the expense of producing sexuals to a stage where raiding by unspecialized, facultative slavemakers, capable of exploiting only small colonies, becomes unlikely owing to a strong work force and (2) Early production of sexual offspring at the cost of colony growth to secure some sexual production in an environment with specialized obligate enslavers, capable of raiding large colonies. We tested the strategies by excavating 30 small to moderately large mounds ofF. podzolica and measured reproductive parameters of colonies in relation to mound size, worker number, and worker size. Mound area predicted worker number satisfactorily. Worker number correlated significantly with worker head width and with number of worker and sexual offspring. With a growing work force, the proportion of sexual offspring increased in the total offspring. Two thirds of the colonies producing sexuals emitted single sex, sex being independent of colony size. Some of the large colonies produced both sexes with a strong bias toward either sex. The unweighted population-level sex ratio did not differ from even, being 0.52 (numerical) or 0.54 (biomass). Very large mounds (not excavated) had small workers and highly male-biased sex ratios, probably owing to energy constraints set by central-place foraging. Population-level colony ontogeny data did not fit either one of the suggested strategies, but imply a mixture of the two. We discuss an alternative, still untested raid-independent explanation to the ontogeny pattern.     

A comparison of mark–release–recapture methods for estimating colony size in the wood ant Formica lugubris

Colony size can be considered the analogue of the body size of a superorganism. Just as body size is important to the physiology of an individual animal, colony size correlates with the life-history and ecology of social insects. Although nest excavation and counting all individuals is the most accurate method for estimating colony size (or nest size), it has the major drawback of being destructive. Alternatively, mark–release–recapture (MRR) can be used repeatedly to measure the size of the same colony or nest. We compared the accuracy and feasibility of four MRR methods and a Mound-Volume method with complete counts from nest excavation for estimating the nest size of F. lugubris, a mound-building wood ant of the Formica rufa group, during the early spring in Scotland. We found that our After-Disturbing method, in which we performed marking and recapturing after gentle disturbance to the top of nest mound, has the best balance between accuracy, non-destructiveness, and time required. We also found that mound volume can be an index of ant nest size under certain conditions. Both non-destructive methods can be used on the same colony or nest repeatedly to monitor nest dynamics.     

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

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

An ant—treehopper mutualism: effects of Formica subsericea on the survival of Vanduzea arquata

Abstract. 1. The numbers of Formica subsericea (Say) were positively correlated with those of the treehopper, Vanduzea arquata Say on branches of black locust.
2. The mean numbers of V.arquata on branches were greater in the second than in the first generation. However, the mean number of ants tending them was significantly lower in the second compared to the first generation.
3. Defoliation of branches resulted in a reduction in the number of mem-bracids on branches and ant numbers declined similarly.
4. Survival of V.arquata was significantly greater when tended by F.subsericea than when ants were exciuded.
5. Ant attendance significantly reduced the number of predators on branches with _Karquata compared to branches lacking both ants and treehoppers. This, in part, explains the greater survival of V.arquata nymphs tended by ants.     

中华红林蚁形态描述、行为观察及社群结构

调查显示,北京百花山上"蚂蚁山"的蚂蚁为我国林区特有的种类——中华红林蚁Formica sinensis。该蚂蚁分布海拔相对较高,通常在针叶林或针阔叶混交林中筑巢,进行社会性生活。成虫有4个品级:大型工蚁、小型工蚁、蚁后和雄蚁。每个成熟蚁巢中有2～4个蚁后,为多蚁后社群结构。夏季蚁巢中同时存在着卵、幼虫、蛹和成虫。目前还只发现工蚁和蚁后休眠越冬。对中华红林蚁的各个虫态和品级进行形态描述和测定,并记述工蚁的觅食、建巢及清巢、防御与进攻等行为习性。首次发现雄蚁具有明显的护巢行为。同时,还发现在工蚁的外出活动中,标记信息素的作用不是很明显。     

Habitat age, breeding system and kinship in the ant Formica fusca

In polygyne ants (multiple queens per colony) factors that affect the distribution and survival of queens may play a key role in shaping the population-wide mating system and colony kin structure. The aim of this paper was to study the breeding system in two populations of different age in the facultatively polygyne ant Formica fusca. Both the observed numbers of queens, and the relatedness patterns among queens, workers and colony fathers were compared in two adjacent populations (ages 17 years and > 100 years) in Southern Finland. The results showed that both the mating system and colony kin structure differed between the study populations. In the old population the relatedness among workers, queens and colony fathers was high. The queens were also related to their mates, resulting in significant inbreeding in workers, but not in queens. Finally, the number of queens per colony fluctuated between years, suggesting queen turnover, and nest-mate queens shared their reproduction unequally (reproductive skew). In the younger population relatedness among queens and workers was lower than in the old population, and the colony fathers were unrelated. Furthermore, inbreeding was absent, and no conclusive evidence was found for reproductive skew among nest-mate queens. Finally, the number of queens per colony appeared more stable between years, although queen turnover occurred also in this population. The observed differences in dispersal and mating behaviour are discussed in the light of a potential connection between population age and habitat saturation.     

The relationship between the distribution of worker sizes and new worker production in the ant Formica neorufibarbis

While it is commonly assumed that variation in worker sizes within a single ant colony increases colony efficiency, there is little causal evidence of a link between worker size variation and colony performance. I tested whether the range of worker sizes within colonies of the ant species Formica neorufibarbis affected new worker production. Removing large workers from colonies lowered the rate of new worker production. A study of unmanipulated colonies indicated that colonies did not maintain a full range of worker sizes; mean worker head widths varied from 0.89-1.24 mm. Colonies naturally missing large workers did not have lower rates of worker production, suggesting that the relative size, not the absolute size, of workers within colonies was important. These are the first results to directly link the range of worker sizes to a component of colony fitness in a natural setting.     

The use of mark-recapture to measure worker number in the rock nesting ant species, Formica neorufibarbis Emery

Summary: I examined the use of a mark-recapture technique to measure colony size and colony growth in the ant species Formica neorufibarbis. I addressed three questions: 1) Is the method reasonably accurate?, 2) is the method precise?, and 3) how many workers does the method kill? I found that estimates of colony sizes based on mark-recapture were similar to those estimated by colony excavation. The error in estimates of worker and cocoon number due to the binomial nature of the mark-recapture method was relatively small, with a mean coefficient of variation of twelve percent for workers and nine percent for cocoons. I estimated that the method killed less than two percent of the workers in a nest.