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''Mycocepurus smithii'' is a species of fungus-growing
ant 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 22,0 ...
from
Latin America Latin America or * french: Amérique Latine, link=no * ht, Amerik Latin, link=no * pt, América Latina, link=no, name=a, sometimes referred to as LatAm is a large cultural region in the Americas where Romance languages — languages derived f ...
. This species is widely distributed geographically and can be found from
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
in the north to
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
in the south, as well as on some
Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean S ...
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 Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and developmen ...
. * 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 Clone or Clones or Cloning or Cloned or The Clone may refer to: Places * Clones, County Fermanagh * Clones, County Monaghan, a town in Ireland Biology * Clone (B-cell), a lymphocyte clone, the massive presence of which may indicate a patholog ...
of the queen. Inside the colony, the ants cultivate a garden of
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
grown with pieces of dead vegetable matter, dead insects, and insect droppings.


Description

Ants of the genus '' Mycocepurus'' are distinctly recognizable for the crown-like cluster of spines on their promesonotum, the fused
mesonotum The mesothorax is the middle of the three segments of the thorax of hexapods, and bears the second pair of legs. Its principal sclerites (exoskeletal plates) are the mesonotum ( dorsal), the mesosternum (ventral), and the mesopleuron (lateral) on ...
and
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum ( dorsal), the prosternum (ventral), and the propleuron ( lateral) on ...
on the front of their
mesosoma The mesosoma is the middle part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the metasoma. It bears the legs, and, in the case of winged insects, the wings. In hymenopterans o ...
or midsection. ''Mycocepurus smithii'' has sharp, protruding propodeal (posterior of the alitrunk) spines unlike ''M. obsoletus'' whose propodeal spines are blunt. Workers also do not have developed promesonotal spines in the center of their crown, which separates ''M. smithii'' from ''M. goeldii'' and similar species.


Reproduction

Initially, ''M. smithii'' was thought to only reproduce asexually because no evidence of male individuals had been found. This led to ''M. smithii'' being recognized as the first fungus-growing ant species to reproduce via thelytokous parthenogenesis, where females, the workers and reproductive queens, are produced asexually. The cytogenetic mechanism of thelytoky is either
apomixis In botany, apomixis is asexual reproduction without fertilization. Its etymology is Greek for "away from" + "mixing". This definition notably does not mention meiosis. Thus "normal asexual reproduction" of plants, such as propagation from cuttin ...
(mitotic parthenogenesis) or
automixis ''Automixis'' is the fusion of (typically haploid) nuclei or gametes derived from the same individual. The term covers several reproductive mechanisms, some of which are parthenogenetic. Diploidy might be restored by the doubling of the chromos ...
with central fusion and low recombination rates. Automixis with central fusion is the
cytogenetic Cytogenetics is essentially a branch of genetics, but is also a part of cell biology/cytology (a subdivision of human anatomy), that is concerned with how the chromosomes relate to cell behaviour, particularly to their behaviour during mitosis a ...
mechanism that was recorded in other asexual ant species. Nests with multiple dealated queens are often found, suggesting that ''M. smithii'' is a
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
species. This appears to be a case of secondary polygyny, and the queens may be daughters of the foundress. A detailed study of many ''M. smithii'' populations across their geographic distribution range (Mexico to Argentina) showed that some ''M. smithii'' populations in the Brazilian
Amazon Amazon most often refers to: * Amazons, a tribe of female warriors in Greek mythology * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon River, in South America * Amazon (company), an American multinational technology co ...
reproduce sexually. This was demonstrated using highly variable
genetic markers A genetic marker is a gene or DNA sequence with a known location on a chromosome that can be used to identify individuals or species. It can be described as a variation (which may arise due to mutation or alteration in the genomic loci) that can be ...
.
Sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, wh ...
was also found stored in the
spermatheca The spermatheca (pronounced plural: spermathecae ), also called receptaculum seminis (plural: receptacula seminis), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, oligochaeta worms and certain other ...
s of queens. Sexual reproduction was suggested as a mechanism for maintaining the
genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. It is dis ...
seen in this species. In summary, ''M. smithii'' is not purely asexual, but instead consists of a "mosaic" of sexual and asexual populations.
Phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
reconstructions and the biology of the species suggest that these sexual populations gave rise to the asexual ones. The mechanism behind the shift to asexuality is still unknown. However,
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention ...
assays and genetic screenings suggest that it is not an
endosymbiont An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
such as ''
Wolbachia ''Wolbachia'' is a genus of intracellular bacteria that infects mainly arthropod species, including a high proportion of insects, and also some nematodes. It is one of the most common parasitic microbes, and is possibly the most common reprodu ...
'' causing the asexuality. In fact, a comparative analysis showed that ''Wolbachia'' endosymbionts do not seem to cause asexuality in ants in general.


Nest architecture

The nests and colonies of ''M. smithii'' were studied in great detail in
Puerto Rico Puerto Rico (; abbreviated PR; tnq, Boriken, ''Borinquen''), officially the Commonwealth of Puerto Rico ( es, link=yes, Estado Libre Asociado de Puerto Rico, lit=Free Associated State of Puerto Rico), is a Caribbean island and unincorporated ...
and Brazil. On the surface, ''M. smithii'' nests can be recognized by their nest mounds consisting of excavated soil and clay. A nest entrance of roughly 1.2 mm in diameter is located in the center of each nest mound. Large ''M. smithii'' nests, which are presumably older, can contain up to 7 or so chambers. Some fungus chambers are shallow whereas others can be found in great depths, as deep as 2 meters. The abandoned chambers are used to deposit waste from the fungus garden and loose soil from chamber construction. The number of nest chambers tends to increase as colonies grow older. Because ''M. smithii'' queens are capable of asexual reproduction, it is believed that colonies can also grow by budding in addition to independent colony foundation. Colonies that grow by budding can result in large colony networks. Workers of ''M. smithii'' ants maintain narrow tunnels (diameter of 1.3 mm), which do not allow two ants to pass each other in the tunnel (head size is around 0.7 mm for workers and 0.9 mm for queens). The tunnels also have a number of slightly larger sections (about 3.6 mm diameter), which would allow passing while also facilitating information exchange. Narrow tunnels are presumably easier (energetically cheaper) to construct and may also aide in leveling the humidity or temperature of the colony or preventing predatory intrusions. In general, ''M. smithii'' colonies are smaller than the colonies of ''M. goeldii''.


Fungal cultivation

When founding a new colony, young queens either shed their wings prior to excavating the nest or just inside. They then excavate a tunnel to a depth of roughly 10 cm (4 in) and create a primary chamber. The dealate, or wingless, queen then carries the wings into the primary chamber and inserts them into the chamber ceiling where the surface of the wings is used as a platform for growing an incipient fungus garden. She will also forage around the nest entrance for caterpillar droppings to feed the fungus garden. The female fore wings of all so-called Paleoattini (the genera ''Mycocepurus'', '' Apterostigma'', and '' Myrmicocrypta'') have a crescent-shaped spot lacking any veins, hairs, and pigmentation, and is thought to provide an "easy to clean" platform for the fungus garden. Queens of the socially parasitic species '' Mycocepurus castrator'' do not found their colonies independently, and the clear spot is absent from their wings. This indirectly supports the idea that the wing spot has a function during the early colony founding and fungus cultivation stage of independently founding ''Mycocepurus'' queens. As the colony matures, workers develop and then tend to the fungus garden, feeding it dried leaves, caterpillar droppings, and other debris from the leaf-litter. One trait of ''M. smithii'' cultivation is that, unlike higher attines, they use a wide diversity of fungal lineages for their gardens. Lineages of ''M. smithii'' have undergone many cultivar shifts over time. This tendency to shift cultivars is hypothesized to be a mechanism for helping to offset some of the costs of asexuality. Also unlike other fungus-growing ants ''M. smithii'' has a
microbiome A microbiome () is the community of microorganisms that can usually be found living together in any given habitat. It was defined more precisely in 1988 by Whipps ''et al.'' as "a characteristic microbial community occupying a reasonably wel ...
that is distinct from the surrounding soil. A Brazilian population of ''M. smithii'' has a fungal cultivar with
gongylidia Gongylidia (singular gongylidium) are hyphal swellings of fungus cultivated by higher-attine genera of fungus-growing ants Fungus-growing ants (tribe Attini) comprise all the known fungus-growing ant species participating in ant–fungus mutu ...
-like structures. This is unusual, because gongylidia are the nutrient rich food bodies produced by the fungi of leaf-cutting ants – and leaf-cutting ants are rather distant relatives of Mycocepurus.


References


External links

*
Photograph of ''Mycocepurus smithii'' at myrmecos.netTaxonomic Information and Synonymy of ''Mycocepurus smithii'' at antcat.orgFurther Information About ''Mycocepurus smithii'' at antwiki.orgFurther Information About the Distribution and Biology of ''Mycocepurus smithii'' at antweb.org
{{Taxonbar, from=Q1954012 Myrmicinae Cloning Insects described in 1893 Hymenoptera of South America Hymenoptera of North America