| Formica truncorum | |
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| Formica truncorum worker | |
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| Species: | F. truncorum
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| Formica truncorum Fabricius, 1804
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Formica truncorum is a species of wood ant from the genus Formica. It is characterized by a grey-brown gaster and bright yellow-red head and thorax, and can be distinguished from other species of Formica by the small erect setae covering its entire body. Workers can range in size from 3.5 to 9.0 mm.[1]
F. truncorum is located in many places worldwide, such as northern Japan, the Jura Mountains and many regions from Italy to Norway. The colonies, which can be polygynous and often polydomous, are spread out in regions of woodland borders, where the ants will make their nests in tree stumps.[2]
Behavior
editEusociality
editF. truncorum is a member of the hymenopteran order of eusocial insects, and so it demonstrates some characteristics specific to eusociality. There is cooperative rearing of young, which is thought to provide a sort of 'life insurance' for the female's brood.[3] If the female were to die during the rearing process, others would take her place to finish raising the brood. The value of this cooperation may justify the cost. Another defining characteristic of eusociality is the evolution of specific castes within the species, some of which are sterile.[3]
Eusocial behavior is thought to have evolved as a result of kin selection within monogamous colonies. In multiply mated colonies, the relatedness between siblings is lowered, which diminishes the benefits of altruistic behavior within the colony. Only in monogamous colonies would the relatedness between individuals in a colony remain high, which could allow the benefits of eusociality to be justified by Hamilton's rule.[3]
Sex ratio conflict
editStudies of F. truncorum have shown that the sex ratio varies between being female-biased or male-biased depending on how many times the queen has mated.[4] In cases where the queen is singly-mated, the relatedness between the workers remains high and the ratio is female-biased. This is thought to occur because of haplodiploidy, which causes the females to be more related to their own sisters than to any potential offspring. This increased relatedness between siblings causes the female workers to skew the sex ratio in their favor. In cases where the queen is multiply mated, this relatedness between siblings is lost, and the queen retains control of the sex ratio, which causes it to be male-biased. These observations also hold true when comparing monogynous and polygynous colonies. Monogynous colonies will have a higher degree of relatedness between the workers, similar to the singly-mated queen, and so will have a female biased sex ratio, while polygynous colonies have more genetic diversity and will be more male-biased.
Resource allocation
editIn addition to sex allocation, the resource allocation of F. truncorum ants to the formation of new reproductives was compared between monogynous and polygynous colonies by measuring the colony’s productivity and mating structure.[5]Monogynous colonies with a high productivity would have excess resources, and they invested these excess resources into the formation of more reproductives. Monogynous colonies typically had a higher ratio of sexuals per adult worker. When they had excess resources, polygynous colonies did not invest in the formation of more reproductives , indicating that the allocation of resources to the formation of reproductives is controlled by the workers.
Effects on fitness
editSingly-mated queens are at a disadvantage to multiply-mated queens who have on average a 37% higher fitness.[6] It is proposed that singly-mated queens remain in the population because the benefits of multiple-mating may have been balanced by the costs of increased predation and risk to disease transmission. The benefits of multiple-mating also decline as it becomes more numerous in the population. Another suggested possibility is that some queens are unable to find a second mate and remain singly-mated.
Dispersal polymorphism
editThe genetic population structure and sociogenetic organization of F. truncorum were compared between monogynous and polygynous colonies by using allele frequency differences between the populations and estimates of relatedness between different subsets of the colony population.[7] The allele frequency differences between subpopulations were significant in the polygynous colonies but not detectable between the subpopulations of the monogynous colonies. Also, the polygynous colonies usually had queens that were related to their male mates, which was not the case in the monogynous colonies, suggesting a difference in dispersal between the two types. Based on this data, polygynous colonies have limited dispersal and will usually mate within the colony, while the monogynous colonies rely on outbreeding and a higher degree of dispersal.
The propensity to disperse also varies between males and females, with the main effect of this being that females that are less prone to dispersal will be more likely to stay in the colony and switch it from monogynous to polygynous.[8] The ability of the female to leave the colony is affect by its physiological condition, where larger females with greater fat stores will preferentially disperse but smaller females are more likely to remain. Male dispersal is also determined by size, and in the case of both sexes, larger males and females are produced in monogynous colonies in accordance with their greater dispersal.
If there is a wide variation in the size of the queens produced, it is likely that monogynous colonies would not remain monogynous for any long period of time, because smaller daughters would remain and form polygynous colonies. To counteract this, queens produced in monogynous colonies must frequently be large. A study of the heritability of queen size shows that there is a significant degree of heritability in size that a daughter can receive from her mother, allowing for monogynous colonies to predictably produce larger queens that will disperse to form independent colonies.[9]
Unicoloniality
editAs seen by comparing the relatedness between the queens and their male mates, polygynous colonies will typically mate intranidally. This results in reproduction that is driven by a very limited dispersal where queens will bud off of the main nest to create a large, polydomous colony. While this normally leads to high relatedness between nestmates, and a high degree of genetic structuring within the colony, nests of F. truncorum in Finland were observed to behave differently from this.[10] These nests will shift sites, depending on the time of year, between a nest for the hibernating season and one for the reproductive season. By marking the nests and using microsatellites, researchers found that the relatedness between nestmates in these colonies was almost zero. This indicates the F. truncorum can form unicolonial populations were the workers migrate between genetically different nests.
Foraging
editThe social divisions of the eusocial ants can allow for the creation of subsets of populations with specific duties assigned to them, which can be applied to a variety of tasks, including foraging. Foraging will usually involve the discovery of small caches of food, with some of them being persistent and reliable and others being very transient. This difference in reliability causes the foragers of F. truncorum to divide into two groups with a specific task.[11] One group will patrol the reliable sites, and the other will scan empty food sites in search of new sources of food. Upon discovering a new source, the foragers will recruit from the nest and will not recruit any foragers already outside. Not all of the recruited foragers will carry food back to the nest, suggesting that some are recruited for defense of the site.
The foraging habits of F. truncorum were also observed by placing baits near foraging areas and then observing the ability of the ants to recruit additional worker to transport the food back to the nest. This process of recruitment is done in a simple manner, where workers that have discovered food will lay a pheromone trail that connects the food to the nest, and additional workers will detect and follow the scent of this trail. Trails that are currently leading towards additional food will have pheromone trails that are more recently placed, and will also likely have a stronger scent due to multiple ants laying down pheromones, which allows the workers to quickly react to changes in the direction of the food source by following the newest and strongest trails. F. truncorum has demonstrated an interested characteristic of its foraging ability where it can successfully follow a trail in light but not in darkness.[12] This suggests a visual component in addition to the pheromone trail that is not fully understood.
Nestmate and kin recognition
editMonogynous colonies and polygynous colonies have shown differences in how they react to nestmate recognition and queen adoption. Monogynous colonies discriminate against, and will not adopt, any non-nestmate female, while polygynous colonies are much more accepting.[13] Monogynous colonies are not totally enclosed, however, and maintain their single-queen status through high female dispersal and low intranidal breeding. While polygynous colonies are more likely to adopt a non-nestmate female, most of the queens still come from adopted daughters within the colony.
Cuticular recognition
editF. truncorum shows an ability to discriminate between nestmates and non-nestmates and also to influence the sex ratio, both of which suggest a phenotypic signal of genetic diversity. A study [14] has indicated that this signal may be the hydrocarbon profile located on the cuticle of each ant. Further examination of these hydrocarbon chains has shown that many contain chains longer than 34 carbons, and while there is enough similarity between cuticles to recognize members of the same colony, they are still diverse enough to be distinct between colonies.[15] Ants which are genetically similar have similar hydrocarbon profiles, which could allow kin to recognize one another through this similarity. Studies have shown that there is a correlation between the cuticular hydrocarbon profile and the dispersal tendencies of particular colonies.[16]
Female preference
editIt was observed in F. truncorum that when two males had their ability to sire offspring compared, one of the two would consistently create more progeny.[17] In spite of this, there was no evidence to support the idea that one male would be particularly better at creating more offspring than the other. This indicates that the female is able to store different amounts of sperm from separate males, which would then result in one male producing significantly more offspring.
References
edit- ^ http://ant.edb.miyakyo-u.ac.jp/E/Taxo/F80703.html
- ^ http://www.antwiki.org/wiki/Formica_truncorum
- ^ a b c Davies, N. B., Krebs, J. R., & West, S. A. (2012). An introduction to behavioral ecology (4th ed.). Oxford: Wiley-Blackwell.
- ^ Sundström, L. (1994). Sex ratio bias, relatedness asymmetry and queen mating frequency in ants. Nature, 367, 266-268. doi:10.1038/367266a0
- ^ Sundstrom, Liselotte. "Sex Allocation and Colony Maintenance in Monogyne and Polygyne Colonies of Formica Truncorum (Hymenoptera: Formicidae): The Impact of Kinship and Mating Structure." The American Naturalist 146.2 (1995): 182-201. Web.
- ^ Sundström, L. and Ratnieks, L. W. Sex ratio conflicts, mating frequency, and queen fitness in the ant Formica truncorum. Behavioral Ecology (1998) 9 (2): 116-121 doi:10.1093/beheco/9.2.116
- ^ Sundstrom, Liselotte. "Genetic Population Structure and Sociogenetic Organisation in Formica Truncorum (Hymenoptera; Formicidae)." Behavioral Ecology and Sociobiology 33.5 (1993): 345-354. Web.
- ^ Sundstrom, Liselotte. "Dispersal Polymorphism and Physiological Condition of Males and Females in the Ant, Formica Truncorum." Behavioral Ecology 6.2 (1995): 132-39. Web.
- ^ Bargum, Katja, Jacobus J. Boomsma, and Liselotte Sundstrom. "A Genetic Component to Size in Queens of the Ant, Formica Truncorum." Behavioral Ecology and Sociobiology 57.1 (2004): 9-16. Web.
- ^ Elias, Marianne, Rainer Rosengren, and Liselotte Sundstrom. "Seasonal Polydomy and Unicoloniality in a Polygynous Population of the Red Wood Ant Formica Truncorum." Behavioral Ecology and Sociobiology 57.4 (2005): 339-49. Web.
- ^ Sundstrom, L. "Foraging Responses OfFormica Truncorum (Hymenoptera; Formicidae); Exploiting Stable vs Spatially and Temporally Variable Resources." Insectes Sociaux 40.2 (1993): 147-61. Web.
- ^ Fortelius, W., and R. Rosengren. "Trail Communication and Directional Recruitment to Food in Red Wood Ants Formica." Annales Zoologici Fennici 24 (1987): 137-46. Web.
- ^ Sundström, L. (1997). Queen acceptance and nestmate recognition in monogyne and polygyne colonies of the ant Formica truncorum. Animal Behaviour, 53(3), 499-510. Retrieved from http://www.sciencedirect.com/science/article/pii/S0003347296903000
- ^ Nielsen, J., Boomsma, J. J., Oldham, N. J., Petersen, H. C., & Morgan, E. D. (1999). Colony-level and season-specific variation in cuticular hydrocarbon profiles of individual workers in the ant Formica truncorm. Insectes Sociaux, 46(1), 58-65. Retrieved from http://dx.doi.org/10.1007/s000400050113
- ^ Akino, Toshiharu. "Cuticular Hydrocarbons of Formica Truncorum (Hymenoptera: Formicidae): Description of New Very Long Chained Hydrocarbon Components." Applied Entomology and Zoology 41.4 (2006): 667-77. Web.
- ^ Johnson, C., and L. Sundstrom. "Cuticular Chemistry of Two Social Forms in a Facultatively Polygyne Ant (Hymenoptera: Formicidae: Formica Truncorum)." Annales Zoologici Fennici 49 (2012): 1-17. Web.
- ^ Keller, Laurent, Liselotte Sundström, and Michel Chapuisat. "Male Reproductive Success: Paternity Contribution to Queens and Workers in Formica Ants."Behavioral Ecology and Sociobiology 41.1 (1997): 11-15. Web.
External links
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