Escovopsis | |
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Escovopsis multiformis SEM images A, B Conidiophores mono- and polycephalous without “swollen cells” C–F Conidiophores mono- and polycephalous with “swollen cells” (red arrows) G, H Vesicles I Phialides J Conidia | |
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Genus: | Escovopsis J.J.Muchovej & Della Lucia (1990) |
Type species | |
Escovopsis weberi J.J.Muchovej & Della Lucia (1990) | |
Species | |
E. aspergilloides Contents | |
Synonyms [2] | |
Escovopsis is a genus of seven formally acknowledged parasitic microfungus species that rely on other fungi to be their hosts. This genus formally circumscribed with a single identified species in 1990; [4] in 2013 three other species were added.
In an early 2015 published study, scientists collected five species of Escovopsis from both genera of fungus-growing ants, Atta and Acromyrmex , four of which came from ant colonies in Brazil and the fifth of which came from Trinidad. These Escovopsis species included E. moelleri , E. microspora , E. weberi , E. lentecrescens , and E. aspergilloides . The research revealed another Escovopsis species ( E. trichodermoides ) isolated and derived from the lower attine ant, Mycocepurus goeldii . It was determined to be a species independent of the other five species because of its complex branch system and varying conidiophores, which lack typical swelling. [5]
A separate study published in January 2015 also looked at the lower attine ant, Mycetophylax morschi, in Brazil and found four strains for the Escovopsis. The scientists that conducted this research also noticed distinguishable features of the Escovopsis such as the type and physicality of the conidiophores and conidiogenous cells and the DNA sequences. They named the species Escovopsis kreiselii . [6]
This genus describes a parasitic fungus capable of horizontally transmitting between colonies, which can affect any of the 47 species of ants commonly known as the leafcutter ants, which are widespread in parts of the Americas. The fungus transmits itself between colonies by utilizing infected external material. [7] Only two species of the pathogenic Escovopsis have been formally made known, although the other rising species and Escovopsis in general is known to act destructively toward to the symbiotic relationship between the ant colonies and their fungus gardens. It was only established as a symbiont of this ant-fungus mutualism proceeding studies that consistently isolated the fungus. [8] However, the leafcutter ants have a defense mechanism against the parasitic fungus, an antifungal actinobacteria from the genus Pseudonocardia .
A study conducted at the University of Costa Rica, San Pedro de Montes de Oca produced results that indicated how strains of Escovopsis in Costa Rica are "better suited" to invade and induce their parasitic effects on the ant colonies (specifically from the Attini tribe and genera Atta and Acromyrmex) than selectively bred fungus. This was concluded from the fact that the Escovopsis strains created several clusters or clades, some more virulent than others, while the fungal cultivar only created a single cluster or clade. This means that the ants in the colonies would be able to reduce, eliminate, and defend against the fungal cultivar much easier and quicker than it would be able to for Escovopsis strains. According to the scientists that undertook the study, the coevolved Escovopsis can be used as a biocontrol agent for the population of these leafcutter ants, which are considered agricultural pests in these areas. [9]
Leafcutter ants communicate through exchanges of chemicals and secrete chemicals made from actinomycete bacteria in order to protect their colonies. The Atta colonies have a hierarchal system of workers and use chemical secretions through metapleural glands, which are capable of producing much phenylacetic acid, an antimicrobial agent. A study demonstrated how the smallest worker ants were able to lower growth rates of Escovopsis spores that were placed into the ants’ fungus garden with this phenylacetic acid. Bioassays showed that Escovopsis from leafcutter ants is not as affected by the acid than that from more basic fungus-growing ants, meaning that there is a need to control the growth of the parasitic Escovopsis fungus because of its virulence as a potential pathogen. [10]
Atta is a genus of New World ants of the subfamily Myrmicinae. It contains at least 17 known species.
Leafcutter ants, a non-generic name, are any of 47 species of leaf-chewing ants belonging to the two genera Atta and Acromyrmex. These species of tropical, fungus-growing ants are all endemic to South and Central America, Mexico, and parts of the southern United States. Leafcutter ants can carry twenty times their body weight and cut and process fresh vegetation to serve as the nutritional substrate for their fungal cultivates.
Fungus-growing ants comprise all the known fungus-growing ant species participating in ant–fungus mutualism. They are known for cutting grasses and leaves, carrying them to their colonies' nests, and using them to grow fungus on which they later feed.
Atta sexdens is a species of leafcutter ant belonging to the tribe Attini, native to America, from the southern United States (Texas) to northern Argentina. They are absent from Chile. They cut leaves to provide a substrate for the fungus farms which are their principal source of food. Their societies are among the most complex found in social insects. A. sexdens is an ecologically important species, but also an agricultural pest. Other Atta species, such as Atta texana, Atta cephalotes and others, have similar behavior and ecology.
The ant–fungus mutualism is a symbiosis seen between certain ant and fungal species, in which ants actively cultivate fungus much like humans farm crops as a food source. There is only evidence of two instances in which this form of agriculture evolved in ants resulting in a dependence on fungi for food. These instances were the attine ants and some ants that are part of the Megalomyrmex genus. In some species, the ants and fungi are dependent on each other for survival. This type of codependency is prevalent among herbivores who rely on plant material for nutrition. The fungus’ ability to convert the plant material into a food source accessible to their host makes them the ideal partner. The leafcutter ant is a well-known example of this symbiosis. Leafcutter ants species can be found in southern South America up to the United States. However, ants are not the only ground-dwelling arthropods which have developed symbioses with fungi. A similar mutualism with fungi is also noted in termites within the subfamily Macrotermitinae which are widely distributed throughout the Old World tropics with the highest diversity in Africa.
Acromyrmex is a genus of New World ants of the subfamily Myrmicinae. This genus is found in South America and parts of Central America and the Caribbean Islands, and contains 33 known species. Commonly known as "leafcutter ants" they comprise one of the two genera of advanced attines within the tribe Attini, along with Atta.
Mycocepurus smithii is a species of fungus-growing ant from Latin America. This species is widely distributed geographically and can be found from Mexico in the north to Argentina in the south, as well as on some Caribbean Islands. It lives in a variety of forested habitats and associated open areas. Two studies published in 2009 demonstrated that some populations of the species consist exclusively of females which reproduce via thelytokous parthenogenesis. A detailed study found evidence of sexual reproduction in some populations in the Brazilian Amazon. Accordingly, M. smithii consists of a mosaic of sexually and asexually reproducing populations. In asexual populations all ants in a single colony are female clones of the queen. Inside the colony, the ants cultivate a garden of fungus grown with pieces of dead vegetable matter, dead insects, and insect droppings.
Pseudonocardia is a genus of the bacteria family Pseudonocardiaceae. Members of this genus have been found living mutualistically on the cuticle of the leafcutter ants because the bacteria has antibiotic properties that protect the fungus grown by the ants. When they are grooming, their legs are passed over their mouth gland that produces the antibiotic and then their legs touch the fungi while they are walking around. The ants have metapleural glands that produce the antimicrobial components to eliminate the Escovopsis fungi. The bacteria may also be found in crypts on the propleural plate. Pseudonocardia is found to have antibiotic properties provided to the leaf-cutter ant to inhibit the growth of Escovopsis, which is a black yeast that parasitizes the leaf-cutter ant. Pseudonocardia can be found in both aquatic and terrestrial ecosystems. Pseudonocardia belongs to the phylum Actinobacteria. Most Actinobacteria grow in soils that are of a neutral pH. Actinobacteria are also important in plant-associated microbial communities are referred to as "free-living." This means that they are not dependent on another organism to live. For example: A non-free-living organism would be a parasite that depends on a host as a food source and a place for shelter. "Free-living" also allows these organisms to require less energy and food for survival. Bacteria from the Pseudonocardia genus are catalase-positive, non-motile, aerobic, non-acid-fast and produce a gram positive reaction. Under the microscope they exhibit branching, rod-shaped organisms.
There are many different strains of Pseudonocardia and a good portion of these strains have been found in China, in soils of the forest, and in Eucalyptus trees of Australia.
Atta vollenweideri, common name chaco leafcutter ant, is a species of leafcutter ant, a New World ant of the subfamily Myrmicinae of the genus Atta. This species is from one of the two genera of advanced attines within the tribe Attini.
Atta mexicana is a species of leaf-cutter ant, a New World ant of the subfamily Myrmicinae of the genus Atta. This species is from one of the two genera of advanced attines within the tribe Attini.
Acromyrmex disciger is a species of leaf-cutter ant, a New World ant of the subfamily Myrmicinae of the genus Acromyrmex. This species is from one of the two genera of advanced attines within the tribe Attini. It is found in the wild naturally in Paraguay.
Acromyrmex fracticornis is a species of leaf-cutter ant, a New World ant of the subfamily Myrmicinae of the genus Acromyrmex. This species is from one of the two genera of advanced attines within the tribe Attini. It is found in the wild naturally in southern Brazil, Paraguay and northern Argentina.
Acromyrmex insinuator is a social parasite of the closely related Acromyrmex echinatior. This specific parasite is of particular interest as it is an opportunity to study the development of social parasitism in the Attini tribe, and provides further evidence for Emery's rule, which theorizes social parasites among insects tend to be parasites of species or genera to which they are closely related to.
Acromyrmex landolti is a species of leaf-cutter ant, a New World ant of the subfamily Myrmicinae of the genus Acromyrmex. This species is from one of the two genera of advanced attines within the tribe Attini.
Acromyrmex laticeps is a species of leaf-cutter ant, a New World ant of the subfamily Myrmicinae of the genus Acromyrmex. This species is from one of the two genera of advanced attines within the tribe Attini.
Acromyrmex striatus is a species of the leaf-cutter ants found in the Neotropics.
Acromyrmex versicolor is known as the desert leafcutter ant. A. versicolor is found during the summer months in the Colorado and Sonoran deserts when there is precipitation. They form large, distinctive nest craters that are covered with leaf fragments. Living and dead leaves are collected by workers and used to cultivate fungus gardens. Each colony can have multiple queens, if they do this is a practice called polygyny, and each queen has her own batch of “starter” fungus. This species does not sting.
Acromyrmex mesopotamicus is a species of leaf-cutter ant, a New World ant of the subfamily Myrmicinae of the genus Acromyrmex. This species is from one of the two genera of advanced attines within the tribe Attini.
Escovopsis aspergilloides is a species of fungus that was rediscovered in 1995 by mycologists Keith A. Seifert, Robert A. Samson and Ignacio Chapela. Escovopsis aspergilloides co-exist in a symbiotic relationship with attini ants - fungus-growing ants. The highly evolved, ancient ant-fungus mutualism has become a model system in the study of symbiosis. In spite of this, the genus Escovopsis was not proposed until 1990 and the first two species were not formally described until the 1990s: E. weberi by Muchovej and Della Lucia in 1990 E. aspergilloides by Seifert, Samson and Chapela in 1995.
Leucoagaricus gongylophorus is a fungus in the family Agaricaceae which is cultivated by certain leafcutter ants. Like other species of fungi cultivated by ants, L. gongylophorus produces gongylidia, nutrient-rich hyphal swellings upon which the ants feed. Production of mushrooms occurs only once ants abandon the nest. L. gongylophorus is farmed by leaf cutter ant species belonging to the genera Atta and Acromyrmex, amongst others.
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