Tandem running

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Two ants walking in tandem. "Follow the leader".jpg
Two ants walking in tandem.
Camponotus consobrinus workers recruit additional nestmates to newly discovered food sources. The lead worker (on the left) has returned to the nest and is leading the remaining workers back to the food source. Shattuck N2-6890-1, Camponotus consobrinus, near Bungendore, NSW.jpg
Camponotus consobrinus workers recruit additional nestmates to newly discovered food sources. The lead worker (on the left) has returned to the nest and is leading the remaining workers back to the food source.

Tandem running is a pair movement coordination observed in ants and termites.

Contents

In ants, tandem running is used for social learning, by which one ant leads another native ant from the nest to the food source it has found. Tandem running is also used to find and choose better, new nest sites to which the colony can emigrate. [1] The follower ant maintains contact with the lead ant by frequently touching the leader's legs and abdomen with its antennae. [2] As predators, scavengers, and herbivores, ants have a variety of food sources, for which they may journey as far as 200 meters from their nest, spraying a scent trail as they go. [3] To lead their kin to new food sources, ants demonstrate one of the few examples of interactive teaching outside of the mammalian class. Social learning by teaching requires that the naive observer change its behavior and acquire some skills or knowledge faster than it would have independently and that the teacher incur some cost. In order for the follower ant to learn landmarks, the leader must travel much slower and make frequent stops to check for his follower. Ultimately, the knowledge of the route to the new food source can be passed throughout the colony as one follower becomes a leader, making tandem running an effective time-saving practice. [4]

In termites, tandem running behavior is used by a mating pair. During a brief period, alates (winged adults) disperse from their nests. Both females and males land on the ground, shed their wings, and run to search for a mating partner. Upon encountering, a pair performs a tandem run, while seeking a suitable site for colony foundation. [5] Tandem running in termites involves communication via sex pheromones, and all mating pairs engage in this process. For these reasons, tandem running in termites is considered a form of sexual behavior.

The following ant frequently touches the legs of the lead ant with its antennae to maintain contact in the tandem run. Tandemrunningdrawing.jpg
The following ant frequently touches the legs of the lead ant with its antennae to maintain contact in the tandem run.

Species that use tandem running

Ants [6] (By genus)

Termites [5] (By genus for each sub/family)

How ants perform tandem running

Individual ants utilize celestial and land cues as well as their own motion detection to navigate, but scout foraging is not their only strategy for finding food resources. Up to 35 percent of ants setting out from the nest forage as tandem pairs. [7]

Studies have found that vision only plays a role in navigation during tandem runs, but plays no role for the followers in following the leader. The follower is only dependent on tactile and phenomenal information. [8] Experiments in which researchers impair the vision of ants found that blind ants were more likely to follow than to lead, but could still occupy either the role of follower or leader, however, for pairs where the leader was fully sighted, their tandem run was usually smoother and faster. Thus, it can be concluded that sight is used for navigation along with other orientation systems, but is not the mechanism by which ants perform their tandem run. [8] Research has shown that leaders rely on tactile feedback from followers during a tandem run. The follower will follow closely behind the leader, and the leader will run rapidly only after it is tapped by the antennae of the follower. [9]

During tandem runs, in addition to tactile signals, ants may also use chemical cues. Pheromone trails help maintain the bond between the ants, facilitate learning, and assist with navigation. [10] The chemical trails may allow followers to stop and examine landmarks before rejoining the leader. Leaders, which are the teachers, are more likely to lay trails during forward tandem runs than during reverse tandem runs. Most leaders deploy their gaster down during forward tandem runs, while the followers do not. During reverse tandem runs, most leaders deploy their gaster in the up or middle position because by this time the ants have learned the route and trails have already been laid, so followers are less likely to get lost. [10]

Follower ants are not necessarily naive foragers. When lead ants were experimentally removed during tandem runs, 40 percent of follower ants successfully reached the foraging area after a brief search, providing evidence that the followers were already experienced foragers. [7]

Costs and benefits

The time it takes for the leader to reach the food source during a tandem run is increased fourfold. [9] However, the follower will be able to find the food source significantly faster during a tandem run compared to searching alone. Additionally, the leader will benefit by having another ant help carry food back to the nest. [9] Experienced foraging ants will often participate in tandem runs as a follower to a familiar tree or foraging site. This may be because trees and other sites are often complex structures to navigate, and the leader may guide the experienced ant to a new food source within the same system. [2] There is also a cost to following ants who get separated from the leader (or "lost") during a tandem run. When tandem running ants were experimentally separated in the lab, lost ants engaged in searching behavior, where they would often return to the point where they were separated from their leader, and their walking speed was slowed by almost 50%. [11] There is a benefit to being separated from the leader in some cases however; independent exploration is critical to the practice of tandem running. Through independent exploration, ants can discover new and more direct routes and thus reduce the duration of their subsequent tandem runs that they will lead. Independent exploration is the basis of route improvement in tandem running. [1]

Teaching and social learning

Tandem running is a form of recruitment and communication that involves teaching and social learning. Experience influences the tendency to engage in tandem running more than age. Temnothorax albipennis of all ages are able to participate in tandem running, but experienced individuals are more likely to do so. Older experienced ants are also more likely to lead slowly and be precise. Young inexperienced ants are capable of leading and following in tandem runs, but they lead faster and are less accurate. [12]

When an individual ant practices on its own it can learn even more. Ants are able to shorten the length of tandem runs and transfer information faster only if they independently explore the area in between being a follower and becoming a leader. [1] Independent exploration is beneficial because it leads to an improved route and more accurate leadership of tandem runs. Only one bout of independent exploration is necessary for improvement, and extra independent explorations do not further improve tandem runs. If a leader happens to lose its follower during a run, it may continue to its destination rather than turning around to find another follower because this is a chance for independent exploration, which can improve future runs. [1] Tandem running combines social learning with individual learning in order to maximize benefits of this practice. The exact information that is transferred between the follower and leader is not known, but the following ant acquires the same information that it would have gotten if it had found the nest on its own. If the new nest site is far from the old one, tandem running allows followers to find it faster than they would have if they used individual exploration. [1]

Tandem running in termites

In Reticulitermes and Coptotermes, female leads the tandem and releases a short-range sex pheromone to guide the male, and the male touches the female's abdomen with its antennae and mouthparts, indicating its continued presence. [13] When a pair is accidentally separated, females pause and males engage in intensive search for the partner. This sexually dimorphic movement facilitates the encounter. [14]

The form of tandem running behavior is variable across termite species. In neoisopteran termites, only females have strong sex-pairing pheromones, and sex role is strictly fixed in heterosexual pairs; females perform leader role, and males perform follower role. On the other hand, sex role is more flexible in other lineages. [5] Also, tandem running behavior can be used between workers in Mastotermes darwiniensis . [15]

In Reticulitermes speratus , tandem running has been observed as homosexual behavior with males following males and females following females in order to reduce predation. By participating in tandem running, the termites increase their encounter risk of predators, but this risk is outweighed by the benefits of decreasing their post-encounter risk. The predator is able to capture only one animal at a time, so by utilizing tandem running, each termite's individual risk of predation is decreased by the dilution effect. [11] During same-sex tandem run in Reticulitermes speratus, one individual expresses the behavior of the other sex. In female-female pairs, follower female shows typical male behaviors. And in male-male pairs, leader males show typical female behavior. [5]

Tandem running is also used for sexual selection in termites. There is often competition between tandem running males over which termite will be in the back position. The male in the back position is thus stronger and larger, and when the pair of tandem runners encounters a female, she will favor the back male. [11]

Related Research Articles

<span class="mw-page-title-main">Ant</span> Family of insects

Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,000 species have been classified. They are easily identified by their geniculate (elbowed) antennae and the distinctive node-like structure that forms their slender waists.

<span class="mw-page-title-main">Termite</span> Social insects related to cockroaches

Termites are a group of detritophagous eusocial insects which consume a variety of decaying plant material, generally in the form of wood, leaf litter, and soil humus. They are distinguished by their moniliform antennae and the soft-bodied and often unpigmented worker caste for which they have been commonly termed "white ants"; however, they are not ants, to which they are only distantly related. About 2,972 extant species are currently described, 2,105 of which are members of the family Termitidae.

<span class="mw-page-title-main">Trophallaxis</span> Transfer of food between members of a community through stomodeal or proctodeal means

Trophallaxis is the transfer of food or other fluids among members of a community through mouth-to-mouth (stomodeal) or anus-to-mouth (proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pheromones, organisms such as symbionts, and information to serve as a form of communication. Trophallaxis is used by some birds, gray wolves, vampire bats, and is most highly developed in eusocial insects such as ants, wasps, bees, and termites.

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

Pogonomyrmex barbatus is a species of harvester ant from the genus Pogonomyrmex. Its common names include red ant and red harvester ant. These large ants prefer arid chaparral habitats and are native to the Southwestern United States. Nests are made underground in exposed areas. Their diets consist primarily of seeds, and they consequently participate in myrmecochory, an ant-plant interaction through which the ants gain nutrients and the plants benefit through seed dispersal. Red harvester ants are often mistaken for fire ants, but are not closely related to any fire ant species, native or introduced.

<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">Blattodea</span> Order of insects that includes cockroaches and termites

Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetic and molecular evidence suggests they evolved from within the cockroach lineage, cladistically making them cockroaches as well. The Blattodea and the mantis are now all considered part of the superorder Dictyoptera. Blattodea includes approximately 4,400 species of cockroach in almost 500 genera, and about 3,000 species of termite in around 300 genera.

<i>Reticulitermes flavipes</i> Species of insect found in North America

Reticulitermes flavipes, the eastern subterranean termite, is the most common termite found in North America. These termites are the most economically important wood destroying insects in the United States and are classified as pests. They feed on cellulose material such as the structural wood in buildings, wooden fixtures, paper, books, and cotton. A mature colony can range from 20,000 workers to as high as 5 million workers and the primary queen of the colony lays 5,000 to 10,000 eggs per year to add to this total.

<i>Eciton burchellii</i> Species of ant

Eciton burchellii is a species of New World army ant in the genus Eciton. This species performs expansive, organized swarm raids that give it the informal name, Eciton army ant. This species displays a high degree of worker polymorphism. Sterile workers are of four discrete size-castes: minors, medias, porters (sub-majors), and soldiers (majors). Soldiers have much larger heads and specialized mandibles for defense. In lieu of underground excavated nests, colonies of E. burchellii form temporary living nests known as bivouacs, which are composed of hanging live worker bodies and which can be disassembled and relocated during colony emigrations. Eciton burchellii colonies cycle between stationary phases and nomadic phases when the colony emigrates nightly. These alternating phases of emigration frequency are governed by coinciding brood developmental stages. Group foraging efforts known as "raids" are maintained by the use of pheromones, can be 200 metres (660 ft) long, and employ up to 200,000 ants. Workers are also adept at making living structures out of their own bodies to improve efficiency of moving as a group across the forest floor while foraging or emigrating. Workers can fill "potholes" in the foraging trail with their own bodies, and can also form living bridges. Numerous antbirds prey on the Eciton burchellii by using their raids as a source of food. In terms of geographical distribution, this species is found in the Amazon jungle and Central America.

<span class="mw-page-title-main">Eusociality</span> Highest level of animal sociality a species can attain

Eusociality is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care, overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society which are sometimes referred to as 'castes'. Eusociality is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform behaviors characteristic of individuals in another caste. Eusocial colonies can be viewed as superorganisms.

<i>Nasutitermes corniger</i> Species of termite

Nasutitermes corniger is a species of arboreal termite that is endemic to the neotropics. It is very closely related to Nasutitermes ephratae. The species has been studied relatively intensively, particularly on Barro Colorado Island, Panama. These studies and others have shown that the termite interacts with many different organisms including a bat that roosts in its nest and various species of ants that cohabit with the termite.

<i>Temnothorax albipennis</i> Species of ant

Temnothorax albipennis, the rock ant is a species of small ant in the subfamily Myrmicinae. It occurs in Europe and builds simple nests in rock crevices.

Trail pheromones are semiochemicals secreted from the body of an individual to affect the behavior of another individual receiving it. Trail pheromones often serve as a multi purpose chemical secretion that leads members of its own species towards a food source, while representing a territorial mark in the form of an allomone to organisms outside of their species. Specifically, trail pheromones are often incorporated with secretions of more than one exocrine gland to produce a higher degree of specificity. Considered one of the primary chemical signaling methods in which many social insects depend on, trail pheromone deposition can be considered one of the main facets to explain the success of social insect communication today. Many species of ants, including those in the genus Crematogaster use trail pheromones.

<i>Reticulitermes</i> Genus of termites

Reticulitermes is a termite genus in the family Rhinotermitidae. They are found in most temperate regions on Earth including much of Asia and the Middle East, Western Europe, and all of North America.

<i>Macrotermes bellicosus</i> Species of insect

Macrotermes bellicosus is a species of Macrotermes. The queens are the largest amongst known termites, measuring about 4.2 inches (110 mm) long when physogastric. The workers average 0.14 in (3.6 mm) in length and soldiers are slightly larger. Bellicosus means "combative" in Latin. The species is a member of a genus indigenous to Africa and South-East Asia.

<i>Temnothorax</i> Genus of ants

Temnothorax is a genus of ants in the subfamily Myrmicinae. It contains more than 380 species.

<i>Temnothorax rugatulus</i> Species of ant

Temnothorax rugatulus is a species of ant in the genus Temnothorax. It is found in North America. More specifically, it is found in the forests of the western United States. Colonies are either monogynous or polygynous. Queens in monogynous colonies are generally larger (marcogynes), about twice the size of conspecific workers; polygynous colonies have smaller queens (microgynes), typically slightly smaller than the workers. T. rugatulus ants are a subdominant ant group.

Nigel R. Franks is an English emeritus professor of Animal Behaviour and Ecology at the University of Bristol. He obtained a BSc and PhD in biology at the University of Leeds. After receiving his BSc in 1977 he began his PhD, during which he spent two years doing field work in Panama on army ants with the Smithsonian Tropical Research Institute. He was awarded the Thomas Henry Huxley Award in 1980 from the Zoological Society of London for the best British PhD in Zoology. He then received a Postdoctoral Fellowship from the Royal Commission for the Great Exhibition of 1851 allowing him to undertake postdoctoral work under Edward O. Wilson at Harvard University before becoming a lecturer at the University of Bath in 1982, later being promoted to full professor in 1995. He moved to the University of Bristol in 2001. He is renowned for his studies of collective animal behaviour, particularly of ant colonies. His Ant Lab at Bristol pioneered the use of Temnothorax as a model ant species for the study of collective decision-making and complex systems. In a 2009 profile in Science he discusses his pioneering use of radio-frequency identification tags (RFID) glued to the backs of each ant for tracking individuals in their society. His book Social evolution in ants with Andrew Bourke was an important contribution to the understanding of kin selection theory and sex ratio theory with respect to social evolution in insects, while his co-authored book Self-organization in biological systems has been cited well over 3000 times

<i>Reticulitermes speratus</i> Species of termite

Reticulitermes speratus, the Japanese termite, is a species of subterranean termite found in Japan, North Korea, and South Korea. It eats decayed wood. It is adapted to withstand the cold temperatures of the temperate regions it inhabits.

<i>Ectatomma opaciventre</i> Species of ant

Ectatomma opaciventre is a South American species of ant from the subfamily Ectatomminae. In the Brazilian savannah, nests occur at low density and in specific microhabitats.

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

References

  1. 1 2 3 4 5 Franklin, Elizabeth L.; Franks, Nigel R. (2012-08-01). "Individual and social learning in tandem-running recruitment by ants". Animal Behaviour. 84 (2): 361–368. doi:10.1016/j.anbehav.2012.05.002. S2CID   53161886.
  2. 1 2 Franks NR, Richardson T (2006). "Teaching in tandem-running ants". Nature . 439 (7073): 153. Bibcode:2006Natur.439..153F. doi: 10.1038/439153a . PMID   16407943. S2CID   4416276.
  3. ↑ Carrol, CR; Janzen, DH (1973). "Ecology of foraging by ants". Annual Review of Ecology and Systematics. 4: 231–257. doi:10.1146/annurev.es.04.110173.001311.
  4. ↑ Davies, N.B., Krebs, J.R., & West, S.A. (2012). An Introduction to Behavioural Ecology. 4th ed, Wiley-Blackwell.
  5. 1 2 3 4 Mizumoto, Nobuaki; Bourguignon, Thomas; Bailey, Nathan W. (2022-11-15). "Ancestral sex-role plasticity facilitates the evolution of same-sex sexual behavior". Proceedings of the National Academy of Sciences. 119 (46): e2212401119. Bibcode:2022PNAS..11912401M. doi: 10.1073/pnas.2212401119 . ISSN   0027-8424. PMC   9674213 . PMID   36346843.
  6. ↑ Hölldobler, Bert; Wilson, Edward O. (1990-01-01). The Ants . Harvard University Press. ISBN   9780674040755.
  7. 1 2 Schultheiss, P., Raderschall, C. A., & Narendra, A. (2015). Follower ants in a tandem pair are not always naïve. Scientific Reports, 5.
  8. 1 2 Franklin, Elizabeth L.; Richardson, Thomas O.; Sendova-Franks, Ana B.; Robinson, Elva J. H.; Franks, Nigel R. (2010-09-10). "Blinkered teaching: tandem running by visually impaired ants" (PDF). Behavioral Ecology and Sociobiology. 65 (4): 569–579. doi:10.1007/s00265-010-1057-2. ISSN   0340-5443. S2CID   46349676.
  9. 1 2 3 Hoppitt, William J. E.; Brown, Gillian R.; Kendal, Rachel; Rendell, Luke; Thornton, Alex; Webster, Mike M.; Laland, Kevin N. (2008-09-01). "Lessons from animal teaching". Trends in Ecology & Evolution. 23 (9): 486–493. doi:10.1016/j.tree.2008.05.008. PMID   18657877.
  10. 1 2 Basari, Norasmah; Laird-Hopkins, Benita C.; Sendova-Franks, Ana B.; Franks, Nigel R. (2014-06-06). "Trail laying during tandem-running recruitment in the ant Temnothorax albipennis" (PDF). Naturwissenschaften. 101 (7): 549–556. Bibcode:2014NW....101..549B. doi:10.1007/s00114-014-1191-1. ISSN   0028-1042. PMID   24903681. S2CID   8422652.
  11. 1 2 3 MATSUURA, KENJI; KUNO, EIZI; NISHIDA, TAKAYOSHI (2002-01-07). "Homosexual Tandem Running as Selfish Herd in Reticulitermes speratus: Novel Antipredatory Behavior in Termites". Journal of Theoretical Biology. 214 (1): 63–70. Bibcode:2002JThBi.214...63M. doi:10.1006/jtbi.2001.2447. PMID   11786032.
  12. ↑ Franklin, E., Robinson, E., Marshall, J., Sendova-Franks, A., Franks, N. (2011). "Do ants need to be old and experienced to teach?" (PDF). Journal of Experimental Biology. 215 (Pt 8): 1287–92. doi: 10.1242/jeb.064618 . PMID   22442366. S2CID   695602 . Retrieved October 15, 2015.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  13. ↑ Mizumoto, Nobuaki; Lee, Sang-Bin; Valentini, Gabriele; Chouvenc, Thomas; Pratt, Stephen C. (2021-07-14). "Coordination of movement via complementary interactions of leaders and followers in termite mating pairs". Proceedings of the Royal Society B: Biological Sciences. 288 (1954): 20210998. doi:10.1098/rspb.2021.0998. ISSN   0962-8452. PMC   8277464 . PMID   34255998.
  14. ↑ Mizumoto, Nobuaki; Dobata, Shigeto (2019-06-07). "Adaptive switch to sexually dimorphic movements by partner-seeking termites". Science Advances. 5 (6): eaau6108. Bibcode:2019SciA....5.6108M. doi:10.1126/sciadv.aau6108. ISSN   2375-2548. PMC   6584256 . PMID   31223644.
  15. ↑ Sillam-Dussès, David; Sémon, Etienne; Lacey, Michael J.; Robert, Alain; Lenz, Michael; Bordereau, Christian (2007-10-16). "Trail-Following Pheromones in Basal Termites, with Special Reference to Mastotermes darwiniensis". Journal of Chemical Ecology. 33 (10): 1960–1977. doi:10.1007/s10886-007-9363-5. ISSN   0098-0331. PMID   17885791. S2CID   22594434.
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