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===Foraging behaviour===
===Foraging behaviour===
The foraging behaviour of wood ants will change according to the environment. Wood ants have been shown to tend and harvest aphids and prey on and compete with other predators for food resources. They tend to prey on the most plentiful members of the community whether they are in the canopies of trees or in the forest foliage. Wood ants seem to favour prey that lives in local canopies near their nest; however, when food resources dwindle, they will seek other trees that are further from the nests and explore more trees instead of exploring the forest floor more thoroughly.<ref>Lenoir, L. "Response of the Foraging Behaviour of Red Wood Ants (''Formica Rufa'' Group) to Exclusion from Trees." Agricultural and Forest Entomology 5.3 (2003): 183-89. Print</ref> This makes foraging for food significantly less efficient, but the rest of the nest will not go out of their way to help the foraging ants.<ref name="Skinner, G. J. 1981">Skinner, G. J., and J. B. Whittaker. "An Experimental Investigation of Inter-Relationships Between the Wood-Ant (''Formica Rufa'') and Some Tree-Canopy Herbivores." Journal of Animal Ecology 50 (1981): 313-26. Print</ref>
The foraging behaviour of wood ants will change according to the environment. Wood ants have been shown to tend and harvest aphids and prey on and compete with other predators for food resources. They tend to prey on the most plentiful members of the community whether they are in the canopies of trees or in the forest foliage. Wood ants seem to favour prey that lives in local canopies near their nest; however, when food resources dwindle, they will seek other trees that are further from the nests and explore more trees instead of exploring the forest floor more thoroughly.<ref>{{cite journal | last1 = Lenoir | first1 = L | year = 2003 | title = Response of the Foraging Behaviour of Red Wood Ants (''Formica Rufa'' Group) to Exclusion from Trees | url = | journal = Agricultural and Forest Entomology | volume = 5 | issue = 3| pages = 183–89 }}</ref> This makes foraging for food significantly less efficient, but the rest of the nest will not go out of their way to help the foraging ants.<ref name="Skinner, G. J. 1981">{{cite journal | last1 = Skinner | first1 = G. J. | last2 = Whittaker | first2 = J. B. | year = 1981 | title = An Experimental Investigation of Inter-Relationships Between the Wood-Ant (''Formica Rufa'') and Some Tree-Canopy Herbivores | url = | journal = Journal of Animal Ecology | volume = 50 | issue = | pages = 313–26 }}</ref>


===Kin behaviour===
===Kin behaviour===
Wood ants have shown aggressive behaviour towards their own species in select situations.<ref>Elton, Charles. "Territory Among Wood Ants (''Formica Rufa'' L.) at Picket Hill." Journal of Animal Ecology 1 (1932): 69-76. Web. <http://www.jstor.org/stable/996></ref><ref name="Skinner, G. J. 1981"/> Intraspecific competition usually occurs early in the spring between workers of competing nests. It has been found that this aggression may be linked to the protection of maintaining territory and trail. By observing skirmishes and trail formation of wood ants, Skinner has noted that the territory surrounding each nest differs between seasons. Permanent foraging trails would be reinforced each season and if another ant from an alien species crossed it, hostile activity would occur. He concludes that most likely, the territory changes based on foraging patterns influenced by seasonal changes.<ref name="Skinner, G. J. 1981"/>
Wood ants have shown aggressive behaviour towards their own species in select situations.<ref name="Skinner, G. J. 1981"/><ref>{{cite journal | last1 = Elton | first1 = Charles | year = 1932 | title = Territory Among Wood Ants (''Formica Rufa'' L.) at Picket Hill | journal = Journal of Animal Ecology | volume = 1 | issue = | pages = 69–76 | jstor=996}}</ref> Intraspecific competition usually occurs early in the spring between workers of competing nests. It has been found that this aggression may be linked to the protection of maintaining territory and trail. By observing skirmishes and trail formation of wood ants, Skinner has noted that the territory surrounding each nest differs between seasons. Permanent foraging trails would be reinforced each season and if another ant from an alien species crossed it, hostile activity would occur. He concludes that most likely, the territory changes based on foraging patterns influenced by seasonal changes.<ref name="Skinner, G. J. 1981"/>


Ants will recognize their nest - mates through chemical signals. Failure in recognition will cause the colony integrity to decay. It has been found that heavy metals accumulated through the environment alter the aggression levels.<ref>Sorvari, Jouni, and Tapio Eeva. "Pollution Diminishes Intra-specific Aggressiveness between Wood Ant Colonies." Science of The Total Environment 408.16 (n.d.): 3189-
Ants will recognize their nest - mates through chemical signals. Failure in recognition will cause the colony integrity to decay. It has been found that heavy metals accumulated through the environment alter the aggression levels.<ref>{{cite journal | last1 = Sorvari | first1 = Jouni | last2 = Eeva | first2 = Tapio | year = | title = Pollution Diminishes Intra-specific Aggressiveness between Wood Ant Colonies | url = http://www.sciencedirect.com/science/article/pii/S0048969710003797# | journal = Science of The Total Environment | volume = 408 | issue = 16| pages = 3189–192 | doi=10.1016/j.scitotenv.2010.04.008}}</ref> This could be due to a variety of factors such as changes in physiological effect or changes in resource levels. The ants in these territories tend to be less productive and efficient. If there was an increased resource competition, one would expect an increase level of aggression, but this is not the case.
192. Web. <http://www.sciencedirect.com/science/article/pii/S0048969710003797#></ref> This could be due to a variety of factors such as changes in physiological effect or changes in resource levels. The ants in these territories tend to be less productive and efficient. If there was an increased resource competition, one would expect an increase level of aggression, but this is not the case.


An experiment was conducted by Moli to perceive how the wood ant diet can affect nest mate recognition. ''Formica rufa'' was housed in a lab under an artificial diet for either seven or 30 days in the presence of homo - colonial queens. After the allotted time, the workers were placed back into their original nests to observe either acceptance or rejection. The wood ants that were kept in the lab for only seven days were recognized; however, the nest mates attacked the wood ants that were kept in the lab for 30 days. Furthermore, a greater degree of aggression was witnessed for those that were heterospecific compared to those that were homospecific. Moli concluded that workers would constantly learn the chemical cues emitted and that the diet of the ants will affect the colony odor and in relation, nest mate identification.<ref>Le Moli, F., and A. Mori. "Field Experiments on Environmental Sources of Nestmate Recognition in Two Species of The group (Hymenoptera Formicidae)." Ethology Ecology & Evolution 1.4 (1989): 329-39. Web. <http://www.tandfonline.com/doi/abs/10.1080/08927014.1989.9525503#.UmoUHJQadvY></ref>
An experiment was conducted by Moli to perceive how the wood ant diet can affect nest mate recognition. ''Formica rufa'' was housed in a lab under an artificial diet for either seven or 30 days in the presence of homo - colonial queens. After the allotted time, the workers were placed back into their original nests to observe either acceptance or rejection. The wood ants that were kept in the lab for only seven days were recognized; however, the nest mates attacked the wood ants that were kept in the lab for 30 days. Furthermore, a greater degree of aggression was witnessed for those that were heterospecific compared to those that were homospecific. Moli concluded that workers would constantly learn the chemical cues emitted and that the diet of the ants will affect the colony odor and in relation, nest mate identification.<ref>{{cite journal | last1 = Le Moli | first1 = F. | last2 = Mori | first2 = A. | year = 1989 | title = Field Experiments on Environmental Sources of Nestmate Recognition in Two Species of The group (Hymenoptera Formicidae) | url = | journal = Ethology Ecology & Evolution | volume = 1 | issue = 4| pages = 329–39 | doi = 10.1080/08927014.1989.9525503 }}</ref>


==Colony structure==
==Colony structure==
Different types of ''Formica rufa'' group species have demonstrated different types of social interaction. Some groups are highly polygynous, with multiple queen colonies forming large networks of connected nests. Others are monogynous, with single-queen colonies.
Different types of ''Formica rufa'' group species have demonstrated different types of social interaction. Some groups are highly polygynous, with multiple queen colonies forming large networks of connected nests. Others are monogynous, with single-queen colonies.
Different ''Formica rufa'' from different regions have been recorded as having traits of being both polygynous and monogynous. The females in the ''Formica rufa'' colonies that are monogynous will separate by flight and establish new nests. Queens in polygynous nests will form new nests in the vicinity of the original nest with the help of workers.<ref>Bernasconi, Christian, Arnaud Maeder, Daniel Cherix, and Pekka Pamilo. "Diversity and Genetic Structure of the Wood Ant Formica Lugubris in Unmanaged Forests." Ann. Zool. Fennici (n.d.): 189-99. Print</ref>
Different ''Formica rufa'' from different regions have been recorded as having traits of being both polygynous and monogynous. The females in the ''Formica rufa'' colonies that are monogynous will separate by flight and establish new nests. Queens in polygynous nests will form new nests in the vicinity of the original nest with the help of workers.<ref>{{cite journal | last1 = Bernasconi | first1 = Christian | last2 = Maeder | first2 = Arnaud | last3 = Cherix | first3 = Daniel | last4 = Pamilo | first4 = Pekka | year = | title = Diversity and Genetic Structure of the Wood Ant Formica Lugubris in Unmanaged Forests | url = | journal = Ann. Zool. Fennici | volume = | issue = | pages = 189–99 }}</ref>
Research has shown that through evolution, polygyny may have arisen through monogyny. One possibility is that monogynous nests due to environmental and physiological conditions may take up new queens.<ref>Goropashnaya, Anna V., Vadim B. Fedorov, and Pekka Pamilo. "Recent Speciation in the ''Formica Rufa'' Group Ants (Hymenoptera, Formicidae): Inference from Mitochondrial DNA Phylogeny." Molecular Phylogenetics and Evolution 32.1 (2004): 198–206. Print</ref><ref>Maeder, Arnaud, Anne Freitag, and Daniel Cherix. "Species- and Nestmate Brood Discrimination in the Sibling Wood Ant Species ''Formica Paralugubris'' and ''Formica Lugubris''." Ann. Zool. Fennici 42 (2005): 201-12. Web</ref>
Research has shown that through evolution, polygyny may have arisen through monogyny. One possibility is that monogynous nests due to environmental and physiological conditions may take up new queens.<ref>{{cite journal | last1 = Goropashnaya | first1 = Anna V. | last2 = Fedorov | first2 = Vadim B. | last3 = Pamilo | first3 = Pekka | year = 2004 | title = Recent Speciation in the ''Formica Rufa'' Group Ants (Hymenoptera, Formicidae): Inference from Mitochondrial DNA Phylogeny | url = | journal = Molecular Phylogenetics and Evolution | volume = 32 | issue = 1| pages = 198–206 }}</ref><ref>{{cite journal | last1 = Maeder | first1 = Arnaud | last2 = Freitag | first2 = Anne | last3 = Cherix | first3 = Daniel | year = 2005 | title = Species- and Nestmate Brood Discrimination in the Sibling Wood Ant Species ''Formica Paralugubris'' and ''Formica Lugubris'' | url = | journal = Ann. Zool. Fennici | volume = 42 | issue = | pages = 201–12 }}</ref>
It has been observed that sometimes in monogynous nests, daughters will be recruited as new reproductive and the nest will become polygynous.<ref>Gyllenstrand, N., P. Seppa, and P. Pamilo. "Genetic Differentiation in Sympatric Wood Ants, ''Formica Rufa'' and ''F. Polyctena''." Insectes Sociaux 51.2 (2004): 139-45. Web. <http://link.springer.com/article/10.1007/s00040-003-0720-2></ref>
It has been observed that sometimes in monogynous nests, daughters will be recruited as new reproductive and the nest will become polygynous.<ref>{{cite journal | last1 = Gyllenstrand | first1 = N. | last2 = Seppa | first2 = P. | last3 = Pamilo | first3 = P. | year = 2004 | title = Genetic Differentiation in Sympatric Wood Ants, ''Formica Rufa'' and ''F. Polyctena'' | url = | journal = Insectes Sociaux | volume = 51 | issue = 2| pages = 139–45 | doi = 10.1007/s00040-003-0720-2 }}</ref>


===Nest splitting===
===Nest splitting===

Revision as of 16:13, 19 June 2015

Formica rufa
Formica rufa worker
Scientific classification
Kingdom:
Animalia
Phylum:
Arthropoda
Class:
Insecta
Order:
Hymenoptera
Family:
Formicidae
Subfamily:
Formicinae
Genus:
Formica
Species:
F. rufa
Binomial name
Formica rufa

Formica rufa, also known as the red wood ant, southern wood ant or horse ant, is a boreal member of the Formica rufa group of ants, and is the type species for that group. It is native to Europe and Anatolia[2] but is also found in North America,[3] in both coniferous and broad-leaf broken woodland and parkland.[3] Workers are bicolored red and brownish-black, with a dorsal dark patch on the head and promensonotum,[3] and are polymorphic, measuring 4.5–9 mm in length.[3] They have large mandibles and like many other ant species they are able to spray formic acid from their abdomens as a defence.[3] Formic acid was first extracted in 1671 by the English naturalist John Ray by distilling a large number of crushed ants of this species.[4]

Description

A caterpillar being bitten by F. rufa.
Patrolling Formica rufa

Nests of these ants are large, conspicuous, dome-shaped mounds of grass, twigs, or conifer needles,[3] often built against a rotting stump, usually situated in woodland clearings where the sun's rays can reach them. Large colonies may have 100,000 to 400,000 workers and 100 queens.[3] Formica rufa is highly polygynous and often re-adopts post-nuptial queens from its own mother colony, leading to old, multi gallery nests which may contain well over a hundred egg-producing females. These colonies can often measure several metres in height and diameter. Formica rufa is aggressively territorial, and will often attack and remove other ant species from the area. Nuptial flights take place during the springtime and are often marked by savage battles between neighbouring colonies as territorial boundaries are re-established.[5] New nests are established by budding from existing nests in the spring,[3] or by the mechanism of temporary social parasitism, the hosts being species of the Formica fusca group, notably F. fusca and F. lemani, although incipient Formica rufa colonies have also been recorded from nests of F. glebaria, F. cunnicularia and similar species including the Lasius genus. A rufa queen will oust the nest's existing queen, lay eggs, and the existing workers will care for her offspring until the nest is taken over.

Diet

A Formica rufa nest
F. rufa nest in meadow near Rila, Bulgaria

The ant's primary diet is aphid honeydew, but they also prey on invertebrates such as insects and arachnids;[3] they are voracious scavengers. Foraging trails may extend 100 m.[3] Larger workers have been observed to forage further away from the nest.[6] Formica rufa is commonly used in forestry and is often introduced into an area as a form of pest management.

Behaviour

Worker behaviour

It has been observed that worker ants in Formica rufa practice parental care or perform cocoon nursing. A worker ant will go through a sensitive phase, where it becomes accustomed to a chemical stimuli emitted by the cocoon. The sensitive phase occurs at an early and specific period. An experiment was conducted by Moli et al. to test how worker ants react to different types of cocoon: homospecific and heterospecific cocoons. If the worker ant is brought up in the absence of cocoons, they will show neither recognition nor nursing behaviour. Both types of cocoons will in fact be opened up by the workers and devoured for nutrients. When accustomed to only the homospecific cocoons, the workers will collect both types of cocoons but only place and protect the homospecific cocoons. The heterospecific cocoons will be neglected and abandoned in the nest and eaten. Lastly, if heterospecific cocoons were injected with extract from the homospecific cocoons, the workers will tend to both types of cocoons equally. This demonstrates that a chemical stimulus from the cocoons seem to be of paramount importance in prompting adoption behaviour in worker ants. However, the specific chemical / stimuli has not been identified.[7]

Foraging behaviour

The foraging behaviour of wood ants will change according to the environment. Wood ants have been shown to tend and harvest aphids and prey on and compete with other predators for food resources. They tend to prey on the most plentiful members of the community whether they are in the canopies of trees or in the forest foliage. Wood ants seem to favour prey that lives in local canopies near their nest; however, when food resources dwindle, they will seek other trees that are further from the nests and explore more trees instead of exploring the forest floor more thoroughly.[8] This makes foraging for food significantly less efficient, but the rest of the nest will not go out of their way to help the foraging ants.[9]

Kin behaviour

Wood ants have shown aggressive behaviour towards their own species in select situations.[9][10] Intraspecific competition usually occurs early in the spring between workers of competing nests. It has been found that this aggression may be linked to the protection of maintaining territory and trail. By observing skirmishes and trail formation of wood ants, Skinner has noted that the territory surrounding each nest differs between seasons. Permanent foraging trails would be reinforced each season and if another ant from an alien species crossed it, hostile activity would occur. He concludes that most likely, the territory changes based on foraging patterns influenced by seasonal changes.[9]

Ants will recognize their nest - mates through chemical signals. Failure in recognition will cause the colony integrity to decay. It has been found that heavy metals accumulated through the environment alter the aggression levels.[11] This could be due to a variety of factors such as changes in physiological effect or changes in resource levels. The ants in these territories tend to be less productive and efficient. If there was an increased resource competition, one would expect an increase level of aggression, but this is not the case.

An experiment was conducted by Moli to perceive how the wood ant diet can affect nest mate recognition. Formica rufa was housed in a lab under an artificial diet for either seven or 30 days in the presence of homo - colonial queens. After the allotted time, the workers were placed back into their original nests to observe either acceptance or rejection. The wood ants that were kept in the lab for only seven days were recognized; however, the nest mates attacked the wood ants that were kept in the lab for 30 days. Furthermore, a greater degree of aggression was witnessed for those that were heterospecific compared to those that were homospecific. Moli concluded that workers would constantly learn the chemical cues emitted and that the diet of the ants will affect the colony odor and in relation, nest mate identification.[12]

Colony structure

Different types of Formica rufa group species have demonstrated different types of social interaction. Some groups are highly polygynous, with multiple queen colonies forming large networks of connected nests. Others are monogynous, with single-queen colonies. Different Formica rufa from different regions have been recorded as having traits of being both polygynous and monogynous. The females in the Formica rufa colonies that are monogynous will separate by flight and establish new nests. Queens in polygynous nests will form new nests in the vicinity of the original nest with the help of workers.[13] Research has shown that through evolution, polygyny may have arisen through monogyny. One possibility is that monogynous nests due to environmental and physiological conditions may take up new queens.[14][15] It has been observed that sometimes in monogynous nests, daughters will be recruited as new reproductive and the nest will become polygynous.[16]

Nest splitting

Wood ants typically have multiple nests so they can move around in case of drastic changes in the environment. This splitting of nests will cause the creating of multiple daughter nests. There are several reasons as to why wood ants would move such as a change in availability of food resources, attack by the population of another colony or a change in the state of the nest itself. During this time, workers, queens and the brood will be transferred from the original nest to the daughter nest in a bilateral direction. The goal is to move to the daughter nest; however, sometimes, the transporting ants may bring an individual back to the original nest. The splitting process can last between a week to over a month long.[17]

Population

Turnover rate of wood ant nests is very quick. Within a period of three years, Dr. Klimetzek counted 248 nests. Furthermore, there was no evidence that a correlation existed between nest age and mortality. It was recorded that smaller nests had lower life expectancy compared to larger nests. The size of the nests would increase as the nest age grew.[18]

Bee paralysis virus

In 2008, the Chronic bee paralysis virus (CBPV) was reported for the first time in this species and another species of ant, Camponotus vagus. CBPV affects bees, ants, and mites.[19]

References

  1. ^ Template:IUCN2006
  2. ^ "Formica rufa".
  3. ^ a b c d e f g h i j Robinson, William H. (2005). Urban Insects and Arachnids: A Handbook of Urban Entomology. Cambridge University Press. p. 247. ISBN 0521812534.
  4. ^ Charles Earle Raven (1986). John Ray, naturalist : his life and works. Cambridge University Press. ISBN 0-521-31083-0.
  5. ^ http://www.bbc.co.uk/naturestop40/ep1/s39.shtml
  6. ^ Wright PJ, Bonser R & Chukwu UO (2000). "The size-distance relationship in the wood ant Formica rufa". Ecological Entomology. 25 (2): 226–233. doi:10.1046/j.1365-2311.2000.00253.x.
  7. ^ Moli, Francesco Le, and Maria Passetti. "Olfactory Learning Phenomena and Cocoon Nursing Behaviour in the Ant Formica Rufa L." Italian Journal of Zoology 45.4 (1978): 389-97. Print
  8. ^ Lenoir, L (2003). "Response of the Foraging Behaviour of Red Wood Ants (Formica Rufa Group) to Exclusion from Trees". Agricultural and Forest Entomology. 5 (3): 183–89.
  9. ^ a b c Skinner, G. J.; Whittaker, J. B. (1981). "An Experimental Investigation of Inter-Relationships Between the Wood-Ant (Formica Rufa) and Some Tree-Canopy Herbivores". Journal of Animal Ecology. 50: 313–26.
  10. ^ Elton, Charles (1932). "Territory Among Wood Ants (Formica Rufa L.) at Picket Hill". Journal of Animal Ecology. 1: 69–76. JSTOR 996.
  11. ^ Sorvari, Jouni; Eeva, Tapio. "Pollution Diminishes Intra-specific Aggressiveness between Wood Ant Colonies". Science of The Total Environment. 408 (16): 3189–192. doi:10.1016/j.scitotenv.2010.04.008.
  12. ^ Le Moli, F.; Mori, A. (1989). "Field Experiments on Environmental Sources of Nestmate Recognition in Two Species of The group (Hymenoptera Formicidae)". Ethology Ecology & Evolution. 1 (4): 329–39. doi:10.1080/08927014.1989.9525503.
  13. ^ Bernasconi, Christian; Maeder, Arnaud; Cherix, Daniel; Pamilo, Pekka. "Diversity and Genetic Structure of the Wood Ant Formica Lugubris in Unmanaged Forests". Ann. Zool. Fennici: 189–99.
  14. ^ Goropashnaya, Anna V.; Fedorov, Vadim B.; Pamilo, Pekka (2004). "Recent Speciation in the Formica Rufa Group Ants (Hymenoptera, Formicidae): Inference from Mitochondrial DNA Phylogeny". Molecular Phylogenetics and Evolution. 32 (1): 198–206.
  15. ^ Maeder, Arnaud; Freitag, Anne; Cherix, Daniel (2005). "Species- and Nestmate Brood Discrimination in the Sibling Wood Ant Species Formica Paralugubris and Formica Lugubris". Ann. Zool. Fennici. 42: 201–12.
  16. ^ Gyllenstrand, N.; Seppa, P.; Pamilo, P. (2004). "Genetic Differentiation in Sympatric Wood Ants, Formica Rufa and F. Polyctena". Insectes Sociaux. 51 (2): 139–45. doi:10.1007/s00040-003-0720-2.
  17. ^ Mabelis, A.A. "Nest Splitting By the Red Wood Ant (Formica Polyctena Foerster)." Netherlands Journal of Zoology 29.1 (1978): 109-25. Web. <http://ingentaconnect.com.libproxy.wustl.edu/content/brill/njz/1978/00000029/00000001/art00005>
  18. ^ Klimetzek, D. "Population Studies on Hill Building Wood-ants of the Formica Rufa -group." Oecologia 48.3 (1981): 418-21. Web. <http://link.springer.com/article/10.1007/BF00346504#page-1>
  19. ^ Detection of Chronic bee paralysis virus (CBPV) genome and its replicative RNA form in various hosts and possible ways of spread.

External links

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