Wikipedia

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, being described already by Linnaeus.[2] It is native to Eurasia, with a recorded distribution stretching from the middle of Scandinavia to the northern Iberia and Anatolia, and from Great Britain to Lake Baikal,[3][2] with unconfirmed reportings of it also to the Russian Far East.[3] There are claims that it can be found in North America,[4] but this is not confirmed in specialised literature,[3] and no recent publication where North American wood ants are listed mentions it as present,[5][3] while records from North America are all listed as dubious or unconfirmed in a record compilation.[6] The workers' heads and thoraces are colored red and the abdomen brownish-black, usually with dark patches on the head and promensonotum,[7] although some individuals may be more uniform reddish and even have some red on the part of the gaster facing the body.[2] In order to separate them from closely related species, specimens needs to be inspected under magnification, where difference in hairiness are among the telling characteristics, with Formica rufa being hairier than per example Formica polyctena but less hairy than Formica lugubris.[2] Workers are polymorphic, measuring 4.5–9 mm in length.[7] 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.[8] Adult wood ants primarily feed on honeydew from aphids. Some groups form large networks of connected nests with multiple queen colonies, while others have single-queen colonies.

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

Description

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

Nests of these ants are large, conspicuous, dome-shaped mounds of grass, twigs, or conifer needles,[4] 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.[4] F. rufa is highly polygynous and often readopts postnuptial queens from its own mother colony, leading to old, multigallery nests that may contain well over 100 egg-producing females. These colonies often may measure several metres in height and diameter. F. rufa is aggressively territorial, and often attacks and removes 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.[9] New nests are established by budding from existing nests in the spring,[4] or by the mechanism of temporary social parasitism, the hosts being species of the F. fusca group, notably F. fusca and F. lemani, although incipient F. rufa colonies have also been recorded from nests of F. glebaria, F. cunnicularia. An F. rufa queen ousts the nest's existing queen, lays eggs, and the existing workers care for her offspring until the nest is taken over.

 
Detail of the head. Picture from antweb.org casent0173863

Diet

edit
 
F. rufa nest
 
F. rufa nest in meadow near Rila, Bulgaria

These ants' primary diet is aphid honeydew, but they also prey on invertebrates such as insects and arachnids;[4] they are voracious scavengers. Foraging trails may extend 100 m.[4] Larger workers have been observed to forage farther away from the nest.[10] F. rufa commonly is used in forestry and often is introduced into an area as a form of pest management.

Behavior

edit

Nursing

edit

Worker ants in F. rufa have been observed to practice parental care or perform cocoon nursing. A worker ant goes through a sensitive phase, where it becomes accustomed to a chemical stimulus 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, it will show neither recognition nor nursing behaviour. Both types of cocoons are opened up by the workers and devoured for nutrients. When accustomed to only the homospecific cocoons, the workers collect both types of cocoons, but only place and protect the homospecific cocoons. The heterospecific cocoons are neglected and abandoned in the nest and eaten. Lastly, if heterospecific cocoons were injected with extract from the homospecific cocoons, the workers tend to both types of cocoons equally. This demonstrates that a chemical stimulus from the cocoons seems to be of paramount importance in prompting adoption behaviour in worker ants. However, the specific chemical / stimulus has not been identified.[11]

Foraging behaviour

edit

The foraging behaviour of wood ants changes 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 seek other trees further from the nests and explore more trees instead of exploring the forest floor more thoroughly.[12] This makes foraging for food significantly less efficient, but the rest of the nest does not help the foraging ants.[13]

Kin behaviour

edit

Wood ants have shown aggressive behaviour toward their own species in certain situations.[13][14] Intraspecific competition usually occurs early in the spring between workers of competing nests. This aggression may be linked to the protection of maintaining territory and trail. By observing skirmishes and trail formation of wood ants, the territory surrounding each nest differs between seasons. Permanent foraging trails are reinforced each season, and if an ant from an alien species crossed it, hostile activity occurs. Most likely, the territory changes based on foraging patterns are influenced by seasonal changes.[13]

Ants recognize their nestmates through chemical signals. Failure in recognition causes the colony integrity to decay. Heavy metals accumulated through the environment alter the aggression levels.[15] 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. Increased resource competition would be expected to increase level of aggression, but this is not the case.

Raiding

edit

Wood ants, particularly those in the Formica species, perform organised and planned attacks on other ant colonies or insects. These planned attacks are motivated by territory expansion, resource acquisition, and brood capture.[16][17] Raids are performed at certain times of the year, when resources may need restocking, and during the day when ants are most active.[17][18] Organised and cooperative strategies for raiding are more specific tactics used by the Formica polyctena species. However, raiding is still an integral behaviour of the Formica rufa group. Scouts will investigate neighbouring nests to raid, marking their targets using pheromones.[19] Wood ants are also capable of counterattack/defending retaliation. Strong defensive measures include guarding entrances to tunnels and having routine patrols of the areas to watch neighbouring nests.[20] Some wood ant species, such as Formica sanguinea, will raid brood, which is then integrated into their colony as workers.[17] This behaviour enables the colony to bolster its workforce without expending energy on raising its brood. The captured brood matures and functions within the raiding colony, helping with foraging and nest maintenance tasks.[17]

Raiding has significant evolutionary and ecological implications. This behaviour can establish dominance hierarchies among colonies and influence the structure of ant communities.[17] Raiding contributes to the success of dominant species by providing access to resources that might otherwise be difficult to obtain.[17] This behaviour also reflects the ants’ ability to adapt their foraging strategies to varying environmental conditions. Wood ants can also alter the distribution of resources in the ecosystem by dominating key food sources.

Resin use

edit

Wood ants intently collect resins from coniferous trees and incorporate them into their nests for various uses. Resin provides wood ants with structural soundness and predator defense to their nests and antimicrobial, antifungal, and pathogen defense when in conjunction with formic acid from their venom gland. [21][22][23]

By leveraging the antimicrobial properties of the resin, wood ants are adequately ensuring and sustaining the health of their colonies.[23][24] Wood ant nests are vulnerable to rapidly spreading microbial loads due to the dense population and organic debris accumulation within large, complex structures. Terpenes and phenolic acids found in coniferous tree resins provide antimicrobial defense and inhibit the growth of pathogens within the nests when mixed with the ants' formic acid.[18][25][26] Nests that have been fortified by resin have significantly less microbial diversity when compared to nests without resin. [23][25][27] By managing their environment, wood ants are proficiently protecting the health of their colonies, with the direct advantages of protecting the queen and developing brood with decreased pathogen exposure. [21][22][23]

Besides antifungal and microbial defense, resin provides value structural integrity to the nest and a protective barrier from potential intruders and predators. Wood ant nests are vulnerable to numerous external threats as they are often large, complex, and above ground. By binding the resin to other organic materials, the nest is provided with cohesive building material, making the nest less prone to collapse.[18]  Incorporating resin also provides nests with waterproofing and weather resistance, another way to prevent fungal growth.[18][28] The stickiness and sometimes toxicity of the resin aid in providing a protective barrier against small arthropods and mites that may attack the nest. Chemically, the resin provides camouflage and deters intruders that may use chemical cues to locate nests.[16][18][21][29]

Colony structure

edit

Polygyny

edit

Polygyny in wood ants (Formica genus) is a colony's social structure that contains multiple reproducing queens. Polygyny may have evolved to enhance colony survival in unstable environments as it allows wood ants to disperse across larger areas by establishing interconnected nests with several queens.[30]This differs from the more commonly observed monogynous social structure of only one reproducing queen within a colony. This behaviour can lead to significant ecological, evolutionary, and colony-level consequences.[30]

Polygyny may have evolved to enhance colony survival in unstable environments as it allows wood ants to disperse across larger areas by establishing interconnected nests with several queens.[31] This differs from a monogamous colony, as a single queen’s reproductive output limits the colony's growth. In a monogamous colony, a new queen will typically leave its nest by flight to find and establish a new nest away from the old one.[32] In a polygynous colony, the new queen will establish its nest nearby, with worker ants helping to connect and create cooperative, large colonies. Polygyny allows for higher genetic diversity within the colony, making the colony less susceptible to pathogens and infections. These polygynous colonies have a more complex social hierarchy and can be more successful in certain ecological contexts because of the combined reproductive efforts of several queens.[33]

Through polygyny, the wood ant colonies exhibit reduced levels of relatedness between workers, which can have negative and positive implications.[33] A negative implication is that there can be reduced cooperation between the ants within a colony. However, this reduced level of cooperation is mitigated by the sheer scale of resources available to polygynous colonies. Besides higher genetic diversity, a positive implication is that the colony has faster growth in numbers due to multiple queens producing broods.[18] With higher numbers, there are more ants to collect resources and carry out raids, but this also has drawbacks. Larger colonies put a lot of structural pressure on the above-ground nest that most wood ants have.[34]

Nest splitting

edit

Wood ants typically have multiple nests so they may relocate in case of drastic changes in the environment. This splitting of nests causes the creation of multiple daughter nests. Several reasons occur as to why wood ants 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 are transferred from the original nest to the daughter nest in a bilateral direction. The goal is to move to the daughter nest, but the transporting ants may bring an individual back to the original nest. The splitting process may last from a week to over a month.[35]

Population

edit

Turnover rate of wood ant nests is very quick. Within a period of three years, Klimetzek counted 248 nests within a 1,640 hectare area under study. Furthermore, no evidence of a correlation between nest age and mortality was found. Smaller nests had lower life expectancy compared to larger nests. The size of the nests increased as the nest aged.[36]

Bee paralysis virus

edit

In 2008, the chronic bee paralysis virus was reported for the first time in this and another species of ants, Camponotus vagus. CBPV affects bees, ants, and mites.[37]

References

edit
  1. ^ Social Insects Specialist Group (1996). "Formica rufa". The IUCN Red List of Threatened Species. 1996: e.T8645A12924924. doi:10.2305/IUCN.UK.1996.RLTS.T8645A12924924.en.
  2. ^ a b c d Seifert, Bernhard (2021). "A taxonomic revision of the Palaearctic members of the Formica rufa group (Hymenoptera: Formicidae) – the famous mound-building red wood ants". Myrmecological News. 31: 133–179. doi:10.25849/myrmecol.news_031:133.
  3. ^ a b c d e Stockan, Jennie A. (2016). Wood Ant Ecology and Conservation. Cambridge University Press. p. 304. ISBN 9781107261402. Archived from the original on 2019-01-11. Retrieved 2022-10-23.
  4. ^ a b c d e f Robinson, William H. (2004). Urban Insects and Arachnids: A Handbook of Urban Entomology. Cambridge University Press. p. 247. ISBN 978-0521812535. Archived from the original on 2022-03-29. Retrieved 2022-11-28.
  5. ^ Jurgensen, Martin F.; Storer, Andrew J.; Risch, Anita C. (2005). "Red wood ants in North America". Ann. Zool. Fennici: 235–242.
  6. ^ "Antmaps.org - Formica rufa". Archived from the original on 2022-10-04. Retrieved 2022-10-23.
  7. ^ a b Collingwood, C.A. (1979). The Formicidae (Hymenoptera) of Fennoscandia and Denmark. Scandinavian Science Press LTD. p. 175. ISBN 978-90-04-27333-7. Archived from the original on 2022-10-23. Retrieved 2022-10-23.
  8. ^ Charles Earle Raven (1986). John Ray, naturalist : his life and works. Cambridge University Press. ISBN 978-0-521-31083-3.
  9. ^ "BBC - Nature's Top 40 - No. 39". Archived from the original on 2015-09-25. Retrieved 2011-06-06.
  10. ^ Wright PJ, Bonser R, Chukwu UO (2000). "The size-distance relationship in the wood ant Formica rufa". Ecological Entomology. 25 (2): 226–233. Bibcode:2000EcoEn..25..226W. doi:10.1046/j.1365-2311.2000.00253.x. S2CID 84140245.
  11. ^ Moli, Francesco Le; Passetti, Maria (1978). "Olfactory Learning Phenomena and Cocoon Nursing Behaviour in the Ant Formica Rufa L." Italian Journal of Zoology. 45 (4): 389–97. doi:10.1080/11250007809440148.
  12. ^ 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. doi:10.1046/j.1461-9563.2003.00176.x. S2CID 84128945.
  13. ^ 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 (1): 313–26. Bibcode:1981JAnEc..50..313S. doi:10.2307/4047. JSTOR 4047.
  14. ^ Elton, Charles (1932). "Territory Among Wood Ants (Formica rufa L.) at Picket Hill". Journal of Animal Ecology. 1 (1): 69–76. Bibcode:1932JAnEc...1...69E. doi:10.2307/996. JSTOR 996.
  15. ^ Sorvari, Jouni; Eeva, Tapio (2010). "Pollution Diminishes Intra-specific Aggressiveness between Wood Ant Colonies". Science of the Total Environment. 408 (16): 3189–192. Bibcode:2010ScTEn.408.3189S. doi:10.1016/j.scitotenv.2010.04.008. PMID 20434195.
  16. ^ a b D'Ettorre, P.; Heinze, J. (2001-04-23). "Sociobiology of slave-making ants". Acta Ethologica. 3 (2): 67–82. doi:10.1007/s102110100038. ISSN 0873-9749.
  17. ^ a b c d e f Mori, Alessandra; Grasso, Donato A.; Le Moli, Francesco (2000). "[No title found]". Journal of Insect Behavior. 13 (3): 421–438. doi:10.1023/A:1007766303588.
  18. ^ a b c d e f DEBOUT, GABRIEL; SCHATZ, BERTRAND; ELIAS, MARIANNE; MCKEY, DOYLE (2007-01-31). "Polydomy in ants: what we know, what we think we know, and what remains to be done". Biological Journal of the Linnean Society. 90 (2): 319–348. doi:10.1111/j.1095-8312.2007.00728.x. ISSN 0024-4066.
  19. ^ Löfqvist, Jan (1976-01-01). "Formic acid and saturated hydrocarbons as alarm pheromones for the ant Formica rufa". Journal of Insect Physiology. 22 (10): 1331–1346. Bibcode:1976JInsP..22.1331L. doi:10.1016/0022-1910(76)90155-4. ISSN 0022-1910.
  20. ^ Adlung, Karl G. (1966-01-12). "A Critical Evaluation of the European Research on Use of Red Wood Ants (Formica rufa Group) for the Protection of Forests against Harmful Insects". Zeitschrift für Angewandte Entomologie. 57 (1–4): 167–189. doi:10.1111/j.1439-0418.1966.tb03822.x. ISSN 0044-2240.
  21. ^ a b c Brütsch, Timothée; Chapuisat, Michel (2014-07-01). "Wood ants protect their brood with tree resin". Animal Behaviour. 93: 157–161. doi:10.1016/j.anbehav.2014.04.024. ISSN 0003-3472.
  22. ^ a b Brütsch, Timothée; Jaffuel, Geoffrey; Vallat, Armelle; Turlings, Ted C. J.; Chapuisat, Michel (April 2017). "Wood ants produce a potent antimicrobial agent by applying formic acid on tree-collected resin". Ecology and Evolution. 7 (7): 2249–2254. Bibcode:2017EcoEv...7.2249B. doi:10.1002/ece3.2834. ISSN 2045-7758.
  23. ^ a b c d Castella, Gregoire; Chapuisat, Michel; Christe, Philippe (2008-04-01). "Prophylaxis with resin in wood ants". Animal Behaviour. 75 (4): 1591–1596. doi:10.1016/j.anbehav.2007.10.014. ISSN 0003-3472.
  24. ^ Castella, Gregoire; Chapuisat, Michel; Christe, Philippe (April 2008). "Prophylaxis with resin in wood ants". Animal Behaviour. 75 (4): 1591–1596. doi:10.1016/j.anbehav.2007.10.014. ISSN 0003-3472.
  25. ^ a b Bos, Nick; Kankaanpää-Kukkonen, Viljami; Freitak, Dalial; Stucki, Dimitri; Sundström, Liselotte (2019-08-26). "Comparison of Twelve Ant Species and Their Susceptibility to Fungal Infection". Insects. 10 (9): 271. doi:10.3390/insects10090271. ISSN 2075-4450. PMID 31454953.
  26. ^ Brütsch, Timothée; Jaffuel, Geoffrey; Vallat, Armelle; Turlings, Ted C. J.; Chapuisat, Michel (2017-03-06). "Wood ants produce a potent antimicrobial agent by applying formic acid on tree-collected resin". Ecology and Evolution. 7 (7): 2249–2254. Bibcode:2017EcoEv...7.2249B. doi:10.1002/ece3.2834. ISSN 2045-7758. PMC 5383563. PMID 28405288.
  27. ^ FEDOROV, VADIM; GOROPASHNAYA, ANNA; JARRELL, GORDON H.; FREDGA, KARL (March 1999). "Phylogeographic structure and mitochondrial DNA variation in true lemmings (Lemmus) from the Eurasian Arctic". Biological Journal of the Linnean Society. 66 (3): 357–371. doi:10.1111/j.1095-8312.1999.tb01896.x. ISSN 0024-4066.
  28. ^ FEDOROV, VADIM; GOROPASHNAYA, ANNA; JARRELL, GORDON H.; FREDGA, KARL (March 1999). "Phylogeographic structure and mitochondrial DNA variation in true lemmings (Lemmus) from the Eurasian Arctic". Biological Journal of the Linnean Society. 66 (3): 357–371. doi:10.1111/j.1095-8312.1999.tb01896.x. ISSN 0024-4066.
  29. ^ de Souza, Danival José; Van Vlaenderen, Johan; Moret, Yannick; Lenoir, Alain (May 2008). "Immune response affects ant trophallactic behaviour". Journal of Insect Physiology. 54 (5): 828–832. Bibcode:2008JInsP..54..828D. doi:10.1016/j.jinsphys.2008.03.001. ISSN 0022-1910. PMID 18430435.
  30. ^ a b Fortelius, Wilhelm (2005). "Mating behaviour in the polygynous/polydomous wood ant Formica aquilonia". Annales Zoologici Fennici. 42 (3): 213–224. ISSN 0003-455X. JSTOR 23735909.
  31. ^ Tsutsui, Neil D.; Suarez, Andrew V.; Holway, David A.; Case, Ted J. (2000-05-16). "Reduced genetic variation and the success of an invasive species". Proceedings of the National Academy of Sciences. 97 (11): 5948–5953. Bibcode:2000PNAS...97.5948T. doi:10.1073/pnas.100110397. ISSN 0027-8424. PMC 18539. PMID 10811892.
  32. ^ Moreau, Corrie S.; Bell, Charles D.; Vila, Roger; Archibald, S. Bruce; Pierce, Naomi E. (2006-04-07). "Phylogeny of the Ants: Diversification in the Age of Angiosperms". Science. 312 (5770): 101–104. Bibcode:2006Sci...312..101M. doi:10.1126/science.1124891. ISSN 0036-8075.
  33. ^ a b Boulay, R.; Arnan, X.; Cerdá, X.; Retana, J. (2014-10-09). "The ecological benefits of larger colony size may promote polygyny in ants". Journal of Evolutionary Biology. 27 (12): 2856–2863. doi:10.1111/jeb.12515. ISSN 1010-061X. PMID 25302869.
  34. ^ Pamilo, Pekka (February 1982). "Genetic population structure in polygynous formica ants". Heredity. 48 (1): 95–106. doi:10.1038/hdy.1982.10. ISSN 0018-067X. PMID 6951823.
  35. ^ Mabelis, A.A. (1978). "Nest Splitting By the Red Wood Ant (Formica polyctena Foerster)". Netherlands Journal of Zoology. 29 (1): 109–25. doi:10.1163/002829679X00124.
  36. ^ Klimetzek, D (1981). "Population Studies on Hill Building Wood-ants of the Formica rufa -group". Oecologia. 48 (3): 418–21. Bibcode:1981Oecol..48..418K. doi:10.1007/BF00346504. PMID 28309762. S2CID 19986018.
  37. ^ Celle, Olivier; Blanchard, Philippe; Olivier, Violaine; Schurr, Frank; Cougoule, Nicolas; Faucon, Jean-Paul; Ribière, Magali (2008). "Detection of Chronic bee paralysis virus (CBPV) genome and its replicative RNA form in various hosts and possible ways of spread" (PDF). Virus Research. 133 (2): 280–284. doi:10.1016/j.virusres.2007.12.011. PMID 18243390. S2CID 16801385.
edit
 
Wikimedia Commons has media related to Formica rufa.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Formica_rufa&oldid=1251982631"