Formica rufa

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Formica rufa
A Formica rufa sideview.jpg
Formica rufa worker
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
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, 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.

Contents

Description

A caterpillar being bitten by F. rufa ForrmicaRufaWithCaterpillar.JPG
A caterpillar being bitten by F. rufa
Patrolling F. rufa Formica rufa on patrol.jpg
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 Formica rufa casent0173863 head 1.jpg
Detail of the head. Picture from antweb.org casent0173863

Diet

F. rufa nest Formica rufa nest 2.jpg
F. rufa nest
F. rufa nest in meadow near Rila, Bulgaria Ant-hill-in-Rila.jpg
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

Nursing

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

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

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

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

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] [27] 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] [28]

Colony structure

Polygyny

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. [29] 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. [29]

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 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. [31] 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. [32]

Through polygyny, the wood ant colonies exhibit reduced levels of relatedness between workers, which can have negative and positive implications. [32] 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. [33]

Nest splitting

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. [34]

Population

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. [35]

Bee paralysis virus

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. [36]

Related Research Articles

<i>Formica exsecta</i> Species of ant

Formica exsecta is a species of ant found from Western Europe to Asia.

<i>Formica</i> Genus of ants

Formica is a genus of ants of the subfamily Formicinae, including species commonly known as wood ants, mound ants, thatching ants, and field ants. Formica is the type genus of the Formicidae, and of the subfamily Formicinae. The type species of genus Formica is the European red wood ant Formica rufa. Ants of this genus tend to be between 4 and 8 mm long. Ants belonging to the Formica genus possess a single knob or bump located between their thorax and abdomen. These ants primarily feed on honeydew, a sugary liquid produced by aphids. Formica ants appear to take on a shepherding role with smaller aphids, relocating them to different parts of plants to ensure a continuous food source for the aphids. By doing so, the ants can establish a relatively sustainable honeydew supply for both themselves and their colony.

<i>Formica rufa</i> species group Group of ants

The Formica rufa group is a subgeneric group within the genus Formica, first proposed by William Morton Wheeler. This group contains the mound-building species of Formica commonly termed "wood ants" or "thatch-mound ants", which build prominent nests consisting of a mound of grass, litter, or conifer needles. The species Formica rufa or the red wood ant is the type species of this subgroup.

<i>Formica cunicularia</i> Species of ant

Formica cunicularia is a species of ant found all over Europe. They are especially common in western Europe and southern England, but they can be found from southern Scandinavia to northern Africa and from Portugal to the Urals. In England, Donisthorpe records the species as having occurred as far north as Bewdley in Worcestershire. In Formica cunicularia, the worker is an ashy grey black color and is usually 4.0–6.5 mm long. The males are found to have a uniformly dark body and are 8.0–9.0 mm long. The queen is yellowish red to dark black and is 7.5–9.0 mm.

<span class="mw-page-title-main">Ant colony</span> Underground lair where ants live, eat, and tend eggs

An ant colony is a population of ants, typically from a single species, capable of maintaining their complete lifecycle. Ant colonies are eusocial, communal, and efficiently organized and are very much like those found in other social Hymenoptera, though the various groups of these developed sociality independently through convergent evolution. The typical colony consists of one or more egg-laying queens, numerous sterile females and, seasonally, many winged sexual males and females. In order to establish new colonies, ants undertake flights that occur at species-characteristic times of the day. Swarms of the winged sexuals depart the nest in search of other nests. The males die shortly thereafter, along with most of the females. A small percentage of the females survive to initiate new nests.

<span class="mw-page-title-main">Carpenter ant</span> Genus of ants (Camponotus spp.)

Carpenter ants are large ants indigenous to many forested parts of the world.

<span class="mw-page-title-main">Pharaoh ant</span> Species of ant

The pharaoh ant is a small (2 mm) yellow or light brown, almost transparent ant notorious for being a major indoor nuisance pest, especially in hospitals. A cryptogenic species, it has now been introduced to virtually every area of the world, including Europe, the Americas, Australasia and Southeast Asia. It is a major pest in the United States, Australia, and Europe. The ant's common name is possibly derived from the mistaken belief that it was one of the Egyptian (pharaonic) plagues.

<span class="mw-page-title-main">Army ant</span> Name used for several ant species

The name army ant (or legionary ant or marabunta) is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limited area.

<span class="mw-page-title-main">Meat ant</span> Common Australian ant

The meat ant, also known as the gravel ant or southern meat ant, is a species of ant endemic to Australia. A member of the genus Iridomyrmex in the subfamily Dolichoderinae, it was described by British entomologist Frederick Smith in 1858. The meat ant is associated with many common names due to its appearance, nest-building behaviour and abundance, of which its specific name, purpureus, refers to its coloured appearance. It is among the best-known species of ant found throughout Australia; it occurs in almost all states and territories except for Tasmania. Its enormous distribution, aggression and ecological importance have made this ant a dominant species.

<i>Formica lugubris</i> Species of ant

Formica lugubris, also known as the hairy wood ant is commonly found in wooded upland areas across northern Eurasia. Colonies construct large thatched mound nests occupied by thousands of workers, and one or more queens. Workers look similar to other species of wood ants, but Formica lugubris workers can be identified by a fringe of hairs that reaches down to their eyes and prominent hairs between the facets of their compound eyes. Workers can reach sizes of up to 9 mm long; queens are larger, reaching 12 mm long.

<i>Formica polyctena</i> Species of ant

Formica polyctena is a species of European red wood ant in the genus Formica and large family Formicidae. The species was first described by Arnold Förster in 1850. The latin species name polyctena is from Greek and literally means 'many cattle', referring to the species' habit of farming aphids for honeydew food. It is found in many European countries. It is a eusocial species, that has a distinct caste system of sterile workers and a very small reproductive caste. The ants have a genetic based cue that allow them to identify which other ants are members of their nest and which are foreign individuals. When facing these types of foreign invaders the F. polyctena has a system to activate an alarm. It can release pheromones which can trigger an alarm response in other nearby ants.

<i>Polyergus breviceps</i> Species of ant

Polyergus breviceps is a species of ant endemic to the United States. It is a social parasite of other ants, namely of Formica gnava but also of Formica occulta and Formica argentea. Polyergus is an inquiline parasite, having lost its ability to take care of its young and themselves. "The workers do not forage for food, feed the young or the queen, or even clean up their own nest". To survive, Polyergus workers raid Formica nests to steal the pupae—which, once hatched, become workers of the mixed nest. This sort of relationship is not unique, of the approximately 8,800 species of ants, at least 200 have evolved some form of symbiotic relationship with one another. What makes Polyergus special is the way a newly mated queen can, all by herself, take over a Formica nest and start a new colony.

<i>Formica sanguinea</i> Species of ant

Formica sanguinea, or blood-red ant, is a species of facultative slave-maker ant in the genus Formica characterized by the ability to secrete formic acid. It ranges from Central and Northern Europe through Russia to Japan, China, the Korean Peninsula, Africa and also the United States. This species is coloured red and black with workers up to 7 mm long.

<i>Formica truncorum</i> Species of ant

Formica truncorum is a species of wood ant from the genus Formica. It is distributed across a variety of locations worldwide, including central Europe and Japan. Workers can range from 3.5 to 9.0mm and are uniquely characterized by small hairs covering their entire bodies. Like all other ants, F. truncorum is eusocial and demonstrates many cooperative behaviors that are unique to its order. Colonies are either monogynous, with one queen, or polygynous, with many queens, and these two types of colonies differ in many characteristics.

<span class="mw-page-title-main">Slave-making ant</span> Species of ants that steal young ants of another species to contribute to their colony

Slave-making ants or slaver ants are brood parasites that capture broods of other ant species to increase the worker force of their colony. After emerging in the slave-maker nest, slave workers work as if they were in their own colony, while parasite workers only concentrate on replenishing the labor force from neighboring host nests, a process called slave raiding.

<span class="mw-page-title-main">Pleometrosis</span> Behaviour in social insects

Pleometrosis is a behavior observed in social insects where colony formation is initiated by multiple queens primarily by the same species of insect. This type of behavior has been mainly studied in ants but also occurs in wasps, bees, and termites. This behavior is of significant interest to scientists particularly in ants and termites because nest formation often happens between queens that are unrelated, ruling out the argument of inclusive fitness as the driving force of pleometrosis. Whereas in other species such as wasps and bees co-founding queens are often related. The majority of species that engage in pleometrosis after the initial stages of colony formation will reduce their colonies number of queens down to one dominant queen and either kill or push out the supernumerary queens. However there are some cases where pleometrosis-formed colonies keep multiple queens for longer than the early stages of colony growth. Multiple queens can help to speed a colony through the early stages of colony growth by producing a larger worker ant population faster which helps to out-compete other colonies in colony-dense areas. However forming colonies with multiple queens can also cause intra-colony competition between the queens possibly lowering the likelihood of survival of a queen in a pleometrotic colony.

<i>Bombus pauloensis</i> Species of bee

Bombus pauloensis is a neotropical bumblebee, formerly known as Bombus atratus, that is found throughout regions of South America, including Colombia, Ecuador, Brazil, and Argentina. It lives in social colonies that include a founder queen/queens, workers and brood. B. pauloensis is somewhat unusual because of its potential to oscillate between polygynous and monogynous nesting cycles. Bombus pauloensis was the first species in the genus Bombus that was discovered to display such polygynous nesting patterns. The polygynous nesting cycles lead to certain specific types of behavior including queen-queen aggression. Nests can also be perennial, which is a characteristic rarely found in other bumblebees. B. pauloensis can be helpful to agricultural because of their ability to pollinate different species of plants. B. pauloensis has been found to occupy a range of geographic areas and climates throughout South America. Colonies have the ability to thermoregulate nests and keep them a little bit warmer than the outside environment. Foraging workers use muscle contractions to maintain stable temperatures and coupe with seasonal and daily fluctuations in temperature.

The social networks of colonies of social insects have been extensively studied as model biological networks.

<span class="mw-page-title-main">Ant supercolony</span> Exceptionally large ant colony

An ant supercolony is an exceptionally large ant colony, consisting of a high number of spatially separated but socially connected nests of a single ant species, spread over a large area without territorial borders. Supercolonies are typically polygynous, containing many egg-laying females. Workers and queens from different nests within the same supercolony can freely move among the nests, and all workers cooperate indiscriminately with each other in collecting food and care of the brood, and show no apparent mutual aggressive behavior.

<i>Formica paralugubris</i> Species of ant

Formica paralugubris is a species of ant. It is a member of the Formica rufa species group native to the Alps in the Palearctic realm. It is a cryptic species, often miscategorized as Formica lugubris due to morphological similarities. The two species are capable of differentiating one another, however; F. paralugubris ants react aggressively towards F. lugubris ants.Formica paralugubris differs in its organization as a supercolony. It has been introduced into North America with the import of pine trees. It does not exhibit characteristics found in other invasive species due to its limited native range.

References

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