Reticulitermes flavipes

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Reticulitermes flavipes
Reticulitermes flavipes K8085-6.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Blattodea
Infraorder: Isoptera
Family: Rhinotermitidae
Genus: Reticulitermes
Species:
R. flavipes
Binomial name
Reticulitermes flavipes
(Kollar, 1837)
Synonyms

Termes flavipes Kollar
Reticulitermes santonensis Feytaud

Contents

Reticulitermes flavipes, the eastern subterranean termite, is the most common termite found in North America. [1] These termites are the most economically important wood destroying insects in the United States and are classified as pests. [1] 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. [1]

Distribution

The eastern subterranean termite (R. flavipes) is the most widely distributed Reticulitermes species. [2] It occurs throughout the eastern United States, including Texas, [3] and can be found as far north as southern Ontario. [4] In 2006, R. flavipes was also recorded in Oregon, [5] on the West coast of the United States.

Reticulitermes santonensis in France is now understood to be a synonym of R. flavipes. [2] The species is now believed to originate in the southeastern United States [6] [7] and to be exotic or invasive in southern Canada, [4] [6] Europe (France, [6] [7] Germany, [6] [8] Italy, [9] the Netherlands [10] ), South America (Uruguay, Chile), [2] [6] and the Bahamas. [11] [6]

Description

Eastern subterranean termites, like other social insects, share resources and divide labor based on a caste system. [1] They live in loosely associated societies called colonies which are composed of both males and females. The termites in the colony are generally organized into the worker caste, the soldier caste, and the reproductive caste. [1]

Worker caste

Worker Reticulitermes flavipes worker.jpg
Worker

Workers are about 3 mm long and are blind, wingless, soft-bodied, creamy white to grayish-white with a round head. [12] They make up the majority of the termites that actually eat the wood. They are sterile and forage for food and water, construct and repair shelter tubes, feed and groom other termites, care for eggs and young, and participate in colony defense. [13]

Soldier caste

Soldier termites are also wingless and resemble workers except that they have a large, rectangular, yellowish-brown head with long black mandibles. The soldiers’ primary function is colony defense [1] and the mandibles are mainly used for crushing enemy ants which may invade the colony. Additionally, R. flavipes has a fontanelle (frontal gland pore) on the forehead from which it secretes a blend of terpenoid compounds. These secretions are synthesised de novo and are also used to defend the colony from predators such as ants. [14] The soldier caste only makes up 1 to 2% of the entire colony. The soldiers are not capable of feeding themselves and rely on the worker termites to provide them with regurgitated food. [1]

Soldiers have been found to protect alates as they begin to leave the nest. Perhaps correspondingly, soldiers populations reach their highest in the Spring as temperatures increase. [15] [16]

Reproductive caste

Wing buds of a nymph Reticulitermes flavipes nymph wing buds.jpg
Wing buds of a nymph

Immature termites on the sexual, or reproductive, line are called nymphs and can be distinguished from workers by the presence of wing buds. Under natural conditions, the work that they do in the nest is negligible compared to that done by workers. They can feed themselves, but there is debate over how long (over how many instars) they retain this ability. [17]

Alates (from the Latin ālātus, "having wings") are winged, sexually mature adults that are capable of dispersal and outbreeding. In R. flavipes, they are 8.5-10mm in length to the tips of their wings [18] and have compound eyes, ocelli, and a dark brown to black fully sclerotized cuticle. [19] As the species name suggests (flāvī, "yellow" + pēs, "foot"), the tarsi are yellowish. [20] To distinguish R. flavipes from similar species (at least in the United States), the position of the ocelli can be used: looking at the head from the side, the distance between the ocellus and the nearest compound eye is at least as large as the diameter of the ocellus. [18] [20]

Alates always develop from nymphs, with the last nymphal stage characterised by particularly long wing buds. After leaving their nests and swarming, male and female alates pair, shed their wings, begin excavating a new nest, and mate. From this point onward, they are called kings and queens, dealates, or primary reproductives. [17]

Wing buds of a brachypterous neotenic Reticulitermes flavipes brachypterous neotenic wing buds.jpg
Wing buds of a brachypterous neotenic

Secondary reproductives, or neotenics, develop from either the reproductive line (brachypterous neotenics) or, in rare cases, the worker line (ergatoids). Ergatoids are the only members of the reproductive caste that have neither wing buds nor wings. Like workers, soldiers, and nymphs, they are eyeless. Ergatoids are seldom found in nature. [17]

Brachypterous neotenics, which are common in the French population of R. flavipes (formerly R. santonensis), and possibly other populations as well, develop from nymphs via a nymph-like pre-brachypterous neotenic stage. [17] They can be distinguished from nymphs by their shorter wing buds (brachy-, "short" + -pterous, "having wings"), darker pigmentation, slight sclerotization, and longer abdomens. Like alates, brachypterous neotenics also have compound eyes and ocelli. Female neotenics lay eggs at a lower rate than queens, but when present in large numbers, they may play a significant role in the growth of the colony. [17]

Life cycle

R. flavipes are opportunistic, and a newly hatched termite can develop into any of a number of castes. At first, it becomes a worker termite and is most likely to remain one for its entire lifespan. Molting can change the worker into a pre-soldier and subsequently, a soldier. The soldier caste is a terminal stage which can no longer molt. [13]

R. flavipes also molts into nymphs, which are the precursors of winged adult termites called alates which are sexually mature. Nymphal termites are non terminal and can revert to the worker stage. These reverted nymphs are called pseudergates. Nymphs and workers can also develop into secondary and tertiary neotenic reproductives respectively. [13]

Behavior

Ventral view of a R. flavipes swarmer Reticulitermes flavipes - swarmer - ventral view, crop.jpg
Ventral view of a R. flavipes swarmer

Because termites are social insects, they share many of their tasks. This can be seen throughout the caste system, where different castes take on different responsibilities for the betterment of the whole colony. R. flavipes cooperate in the rearing of young and also share their resources with the nest. [13]

Swarming is the sudden, dramatic appearance of R. flavipes alates in the daytime from February to April. After this behavior male and female alates lose their wings, pair up, and form new colonies. [13]

R. flavipes is mobile throughout its life and no permanent central nest area exists. Therefore, all termite castes can be found in any of the different sites occupied by a colony. Their activity is determined by food, moisture and temperature, and movement is usually driven by one of these necessities. Termites feed on anything made of or containing cellulose, but can tunnel into non-cellulose containing material to gain access to their destination. This behavior can be destructive to human activities. [13]

Gut microbiota

Flagellates

Discovery

The first person to observe and formally describe symbiotic flagellates in R. flavipes was the American paleontologist Joseph Leidy. In 1877, he reported his findings to the Academy of Natural Sciences of Philadelphia, describing three new species which he mistakenly took to be parasitic ciliates: Trichonympha agilis, Pyrsonympha vertens, and Dinenympha gracilis. [21]

Prof. LEIDY remarked that in seeking small animals beneath stones and fragments of wood in our forests, observing the very common White Ant, Termes flavipes, he noticed that the intestine of the insect, seen in the translucent abdomen, was distended with brown matter. Feeling curious to learn the exact nature of this matter, he was surprised to find that it consisted largely of infusorial and other parasites, mingled with minute particles of decayed wood. In many instances the parasites are so numerous as to make up the greater portion of the bulk of the intestinal pulp. Every individual he had examined, of workers, soldiers, and winged forms, was infested with the parasites, which may be estimated by millions.

— Proceedings of the Academy of Natural Sciences of Philadelphia (1877) [21]

Species present in Reticulitermes flavipes

Flagellates from a Reticulitermes flavipes gut

So far, least 9 genera and 11 species of symbiotic flagellate have been found in the R. flavipes hindgut: [22] [23]

A 2003 study that focused on oxymonad flagellates found evidence of at least one additional undescribed Dinenympha species. They also found what they believe is an undescribed "second" Pyrsonympha species in addition to P. vertens . The authors question whether P. major , which was first described in Reticulitermes hesperus, is truly found in R. flavipes and consider it a case of mistaken identity. [24]

Role in digestion

Flagellates are a feature of all termite families except Termitidae, the so-called "higher termites". They are found exclusively in the hindgut, especially the paunch, an enlarged section of the hindgut with an anaerobic interior that serves as a fermenter. In R. flavipes, wood eaten by the termite is first broken up with the mandibles, treated with host endoglucanases from the salivary glands, [25] ground up into small particles in the gizzard, and then treated with additional host cellulases in the midgut, freeing glucose for immediate absorption. It then passes into the paunch, where flagellates take up the partly-digested wood particles through endocytosis. [26] The flagellates break lignocellulose (xylan or cellulose) down into simple sugars, which they ferment for their own energy needs, producing CO2, H2, and short-chain fatty acids such as acetate as waste products. The short-chain fatty acids (which are also produced by homoacetogenic gut bacteria from CO2 and H2) can be directly absorbed by the termite host. [26] [27] The vast majority of digestion takes place in the paunch: almost all xylanase activity, just over one quarter of the endoglucanase activity, and almost two thirds of the exoglucanase activity in the R. flavipes digestive system is localised in the hindgut and attributed to gut flagellates. [25]

Although this is an obligate symbiosis, newly hatched larvae do not have gut flagellates. These are acquired when the larvae are fed with flagellate-containing anal fluids from other members of the colony (proctodeal trophallaxis). Proctodeal trophallaxis is also used to replenish flagellates and other gut symbionts after each moult. [26]

Bacteria

Spirochaetes and flagellates from a R. flavipes gut Unidentified spirochaetes from a Reticulitermes flavipes gut.jpg
Spirochaetes and flagellates from a R. flavipes gut

Most bacterial species in the termite gut are difficult or impossible to culture, [26] [28] so methods like 16S ribosomal RNA sequencing are used to identify which groups are present. One such analysis of R. flavipes worker guts uncovered representatives of the Endomicrobia (Elusimicrobiota), Actinomycetota, Pseudomonadota, Bacillota, Bacteroidota, and Spirochaetota. [28] The dominant bacterial taxon in the core gut microbiome is the genus Treponema (Spirochaetota), which accounted for approximately 32% of sequences in another 16S rRNA study. [29] The same study identified Endomicrobia (phylum:Elusimicrobiota), which are predominant endosymbionts several protozoa in the termtie gut. [29] Artificial defaunation of Reticulitermes species by force-feeding on starch or starvation leads to a loss of flagellates and, by association, these endosymbiotic Endomicrobia, and an increase in abundance of free-living relatives. [30] A rare free-living member of this class, Endomicrobium proavitum , the first Endomicrobia species to be cultured and named, was isolated from sterile-filtered gut homogenates from defaunated (starch-fed) R. flavipes workers. It is believed to play a role in nitrogen fixation. [31]

While some gut bacteria live free in the gut lumen or attached to the gut wall, many others live in close association with flagellates. These include uncultured members of Endomicrobia (Elusimicrobiota) and other taxa that live as cytoplasmic symbionts, as well as epibiotic symbionts that attach to the outer surface of cells. In R. flavipes, cytoplasmic symbionts have been identified in flagellates such as Trichonympha agilis (Hypermastigida) and Pyrsonympha vertens (Oxymonadida). [32] The latter is also characterised by spirochaete epibiotic symbionts. [33] In other termite species, members of the spirochaete genus Treponema have been identified as endo- [34] and ectosymbionts [35] of flagellates: similar associations may explain Treponema's dominance in the R. flavipes gut. [29]

Disruption of the gut microbiota using agents like antibiotics [36] or boric acid [37] , a common agent used in preventative treatment, have been shown to cause irrecoverable damage to digestion and lead to the rise of opportunistic pathogens.

Archaea

Archaea isolated from the R. flavipes hindgut include the H2- and CO2-utilizing methanogens Methanobrevibacter cuticularis and curvatus . [38] These two species are believed to be the dominant methanogens in R. flavipes and are found in the peripheral, microxic region of the hindgut, on or in close proximity to the gut wall, where they are sometimes attached to filamentous prokaryotes. [38]

Caste-specific differences

Although there is a core microbiome shared among castes and colonies, the alate caste gut is characterised by a significantly lower abundances of bacteria in the genus Treponema (Spirochaetota) and the class Endomicrobia (Elusimicrobia) compared to workers and soldiers. [29] Flagellates in the class Parabasalia and the order Oxymonadida are also significantly less abundant in the alate gut, with the latter significantly reduced in dealates as well. [29]

Human impact

Along with Reticulitermes virginica, R. flavipes is responsible for 80% of the $2.2 billion spent annually in the United States on termite control. [39] Termites feed on wood cellulose, meaning that their presence in human made structures often goes unnoticed for lengthy periods of time. A termite infested timber will appear structurally sound from the outside, while inside it will have a honeycombed appearance. To detect the presence of R. flavipes the observer can test the integrity of the wood by tapping it with a screwdriver. If present, R. flavipes is found at, near or below ground level. Trim work, sub flooring, flooring, and the structural timbers are the areas of a building most susceptible to termite damage. The Eastern subterranean termite is considered a serious economic timber pest and it is estimated that in high activity areas more than 1 in 5 homes have been or will be attacked. [40]

Termite control methods include: physical barriers, chemical treatments, and physical treatments (such as heat, freezing, electrocution and microwave irradiation). [41]

Structural damage to buildings is not R. flavipes’ only impact on humans. Termites also play a critical role in the decomposition of organic matter in natural communities. Without termites, the accumulation of dead organic matter on the forest floor would become detrimental to integrity of that forest. The benefits provided by R. flavipes in terms of their contribution to environmental regulation may far outweigh the disadvantages they pose. [42]

See also

Related Research Articles

<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">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>Trichonympha</i> Genus of flagellated protists

Trichonympha is a genus of single-celled, anaerobic parabasalids of the order Hypermastigia that is found exclusively in the hindgut of lower termites and wood roaches. Trichonympha’s bell shape and thousands of flagella make it an easily recognizable cell. The symbiosis between lower termites/wood roaches and Trichonympha is highly beneficial to both parties: Trichonympha helps its host digest cellulose and in return receives a constant supply of food and shelter. Trichonympha also has a variety of bacterial symbionts that are involved in sugar metabolism and nitrogen fixation.

The Sentricon Termite Colony Elimination System is a subterranean termite pest control product developed and manufactured by Corteva. It was introduced in 1995 as a termite baiting system and an alternative to liquid termicide soil barriers. It eliminates all members of the termite colony, including those of the Formosan subterranean termite colonies.

<span class="mw-page-title-main">Macrotermitinae</span> Subfamily of termites

The Macrotermitinae, the fungus-growing termites, constitute a subfamily of the family Termitidae that is only found within the Old World tropics.

<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.

<span class="mw-page-title-main">Kalotermitidae</span> Family of termites

Kalotermitidae is a family of termites, commonly known as drywood termites. Kalotermitidae includes 21 genera and 419 species. The family has a cosmopolitan circumtropical distribution, and is found in functionally arid environments.

Pyrsonympha is a genus of Excavata.

<i>Zootermopsis angusticollis</i> Species of termite

Zootermopsis angusticollis is a species of termite (Isoptera) in the family Archotermopsidae, a group known as the Pacific dampwood termites, or the rottenwood termites. As their name suggests, the dampwood termites can only survive by living off of wood that contains high amounts of moisture. They are found along the wet environments of the Pacific coast of North America. Most are found in the states of California, Oregon, Washington, Idaho, Western Nevada and in southern British Columbia. Termites are well known to be destroyers of wood, and although the dampwood termites can cause some damage, they are not as notoriously known to cause as much damage to buildings as the drywood termites. They occasionally have been carried to other parts of the country through wood shipments, but have not been able to become established in these areas due to undesirable environmental conditions.

<i>Coptotermes gestroi</i> Species of termite

Coptotermes gestroi, commonly known as the Asian subterranean termite is a small species of termite that lives underground. Both this species and the Formosan subterranean termite are destructive pests native to Asia, but have spread to other parts of the world including the United States. In Asia, this species is known as the Philippine milk termite.

<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>Incisitermes minor</i> Species of termite

Incisitermes minor is a species of termite in the family Kalotermitidae known commonly as the western drywood termite. It is native to western North America, including the western United States and northern Mexico. It has been found in many other parts of the United States, all the way to the East Coast. It has been reported from Toronto. It has been introduced to Hawaii. It has been noted in China and it is not uncommon in Japan. This is an economically important pest of wooden structures, including houses. In California and Arizona alone its economic impact is estimated to be about $250 million per year.

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

Reticulitermes hesperus, the western subterranean termite, is a species of termite in the family Rhinotermitidae. It is found in Central America and North America. R. hesperus is native to the coast between British Columbia and Southern California. Like other subterranean termites, they live underground, where they have elaborate eusocial societies composed of a queen, workers, and soldiers, as well as a rotating case of sexually reproductive adults and their larval and immature offspring. The reproductive adults are the only ones with functional wings. The reproductive adults will swarm on warm days in spring and fall, particularly after a rain event, looking for mating partners. These termites prefer moist living environments and prefer to consume wood that has already been partially decayed by saprotrophic fungus.

Prorhinotermes simplex, the Cuban subterranean termite, is a species of lower termite in the genus Prorhinotermes. It is found in Colombia. Like others in its genus, it is a single-site nesting termite that moves to a new food source when theirs is gone, and it lacks a true worker caste.

<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.

Reticulitermes arenincola is a species of subterranean termite native to North America.

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

Reticulitermes virginicus is a species of subterranean termite native to North America, found often in the southern United States. It was described in 1907.

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

Reticulitermes tibialis, the arid-land subterranean termite, is a species of termite in the family Rhinotermitidae. It is found in the United States, mostly in the western half, occurring in deserts, prairies and other dry locations.

<i>Heterotermes</i> Genus of termites

Heterotermes is a genus of subterranean termites belonging to the family Rhinotermitidae. The genus has an almost cosmopolitan distribution. One of their closest relatives is the Reticulitermes genus.

<i>Heterotermes aureus</i> Species of termite

Heterotermes aureus, commonly known as the desert subterranean termite, is a species of termite in the family Rhinotermitidae. It is native to the deserts of North America where the colony has an underground nest.

References

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  12. ↑ dBugger.org. "Subterranean Termite Identification & Castes" . Retrieved 2013-07-26.
  13. 1 2 3 4 5 6 Susan C. Jones. "Termite Control". Ohio State University . Retrieved February 20, 2009.
  14. ↑ Lori J. Nelson; Laurence G. Cool; Brian T. Forschler; Michael I. Haverty (2001). "Correspondence of Soldier Defense Secretion Mixtures with Cuticular Hydrocarbon Phenotypes for Chemotaxonomy of the Termite Genus Reticulitermes in North America". Journal of Chemical Ecology . 27 (7): 1449–1479. Bibcode:2001JCEco..27.1449N. doi:10.1023/A:1010325511844. PMID   11504039. S2CID   9781034.
  15. ↑ Haverty, M. I.; Howard, R. W. (March 1981). "Production of soldiers and maintenance of soldier proportions by laboratory experimental groups of Reticulitermes flavipes (Kollar) and Reticulitermes virginicus (Banks) (Isoptera:Rhinotermitidae)". Insectes Sociaux. 28 (1): 32–39. doi:10.1007/BF02223620. ISSN   0020-1812. S2CID   20011514.
  16. ↑ Tian, Li; Zhou, Xuguo (2014). "The Soldiers in Societies: Defense, Regulation, and Evolution". International Journal of Biological Sciences. 10 (3): 296–308. doi:10.7150/ijbs.6847. ISSN   1449-2288. PMC   3957085 . PMID   24644427.
  17. 1 2 3 4 5 Lainé, L.V.; Wright, D.J. (2003). "The life cycle of Reticulitermes spp. (Isoptera: Rhinotermitidae): what do we know?". Bulletin of Entomological Research. 93 (4): 267–278. doi:10.1079/BER2003238. PMID   12908912.
  18. 1 2 Scheffrahn, Rudolf H.; Su, Nan-Yao (December 1994). "Keys to Soldier and Winged Adult Termites (Isoptera) of Florida". The Florida Entomologist. 77 (4): 460. doi: 10.2307/3495700 . JSTOR   3495700.
  19. ↑ Jaques, Roger G. Bland, H.E. (2010). How to know the Insects (3rd ed.). Long Grove, IL.: Waveland Press. p. 116. ISBN   978-1577666844.{{cite book}}: CS1 maint: multiple names: authors list (link)
  20. 1 2 Messenger, Matthew T. (2004). "The Termite Species of Louisiana: An Identification Guide". New Orleans Mosquito & Termite Control Board Bulletin (4–04, 3rd Edition).
  21. 1 2 Leidy, Joseph (1877). "On Intestinal Parasites of Termes flavipes". Proceedings of the Academy of Natural Sciences of Philadelphia. 29: 146–149. JSTOR   4060180.
  22. ↑ Lewis, J. L.; Forschler, B. T. (1 November 2006). "A Nondichotomous Key to Protist Species Identification of Reticulitermes (Isoptera: Rhinotermitidae)". Annals of the Entomological Society of America. 99 (6): 1028–1033. doi: 10.1603/0013-8746(2006)99[1028:anktps]2.0.co;2 .
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  24. ↑ Stingl, Ulrich; Brune, Andreas (April 2003). "Phylogenetic Diversity and Whole-Cell Hybridization of Oxymonad Flagellates from the Hindgut of the Wood-Feeding Lower Termite Reticulitermes flavipes". Protist. 154 (1): 147–155. doi:10.1078/143446103764928530. PMID   12812374.
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