Myrmecopterula Moniliformis 2
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''Myrmecopterula'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Pterulaceae The ''Pterulaceae'' are a family (biology), family of fungi in the order Agaricales. According to a 2008 estimate, the family contained 99 species previously distributed among 12 genus, genera. More recent data from molecular phylogenetic reconst ...
.
Basidiocarps In fungi, a basidiocarp, basidiome, or basidioma () is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do no ...
are
clavarioid The clavarioid fungi are a group of fungi in the Basidiomycota typically having erect, simple or branched basidiocarps (fruit bodies) that are formed on the ground, on decaying vegetation, or on dead wood. They are colloquially called club fungi a ...
and are associated with
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
domestication by members of the genus ''
Apterostigma ''Apterostigma'' is a genus of New World ants of the subfamily Myrmicinae. Two species have been described from fossils preserved in Dominican amber, while the others are extant. They are fungus-growing ants, though, unlike the majority of othe ...
''.


Taxonomy

''Myrmecopterula'' was created in 2020 when the ''Pterulaceae'' was reclassified based on
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analysis and split into ''
Pterula ''Pterula'' is a genus of fungi in the '' Pterulaceae'' family. The genus has a widespread distribution, especially in tropical regions, and contains about 50 species. One such species, ''Pterula sp. 82168'', has yielded potential antifungal anti ...
'', ''Myrmecopterula'', ''
Pterulicium ''Pterulicium'' is a genus of fungi in the Pterulaceae family. The genus was previously monotypic, containing the single species ''Pterulicium xylogenum'', found in southeast Asia. However in 2020 a major reclassification of the Pterulaceae fam ...
''. and ''
Phaeopterula ''Phaeopterula'' is a genus of fungi in the family ''Pterulaceae''. Basidiocarps are clavarioid and resemble species of ''Pterula''. Taxonomy This genus was originally described by the German mycologist Paul Christoph Hennings in 1905 but most ...
''. ''Myrmecopterula'' is described as differing from ''Pterula'' in the presence of a cotton-like subiculum. In the context of
mycology Mycology is the branch of biology concerned with the study of fungus, fungi, including their Taxonomy (biology), taxonomy, genetics, biochemistry, biochemical properties, and ethnomycology, use by humans. Fungi can be a source of tinder, Edible ...
the subiculum is defined as a net, cottony or crust like growth of
mycelium Mycelium (: mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Its normal form is that of branched, slender, entangled, anastomosing, hyaline threads. Fungal colonies composed of mycelium are fo ...
from which hyphae or fruiting bodies are produced. The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
, ''M. moniliformis'' was originally classified as ''Lachnocladium moniliforme'' by the German mycologist
Paul Christoph Hennings Paul Christoph Hennings (November 27, 1841 – October 14, 1908) was a German mycologist and herbarium curator. He discovered the study of cryptogams and mushrooms as a volunteer at the botanical garden. Although circumstances initially prevente ...
in 1904. It was classified as ''Thelephora clavarioides'' by the Portuguese mycologist
Camille Torrend Camille Torrend (1875-1961) was a Portuguese clergyman and mycologist. He was active in France, Portugal, Ireland and Brazil. He was a professor of botany and phytopathology at the Imperial Agricultural School of Bahia. Torrend edited the exsiccat ...
in 1914. It was reclassified as ''Pterula moniliformis'' by the English mycologist
Edred John Henry Corner Edred John Henry Corner FRS (12 January 1906 – 14 September 1996) was an English mycologist and botanist who occupied the posts of assistant director at the Singapore Botanic Gardens (1929–1946) and Professor of Tropical Botany at the Un ...
in 1952. ''M. nudihortorum'' and ''M. velohortorum'' were originally classified as ''
Pterula ''Pterula'' is a genus of fungi in the '' Pterulaceae'' family. The genus has a widespread distribution, especially in tropical regions, and contains about 50 species. One such species, ''Pterula sp. 82168'', has yielded potential antifungal anti ...
'' species by the American mycologist
Bryn Tjader Mason Dentinger Bryn is a Welsh word meaning hill. It may also refer to: Places United Kingdom England * Bryn, Cheshire, a location * Bryn, Greater Manchester ** Bryn (ward), an electoral ward in Wigan ** Bryn railway station * Bryn, Shropshire, a location ...
in 2014. These species were ultimately all reclassified as ''Myrmecopterula'' by the mycologists Caio A. Leal-Dutra,
Bryn Tjader Mason Dentinger Bryn is a Welsh word meaning hill. It may also refer to: Places United Kingdom England * Bryn, Cheshire, a location * Bryn, Greater Manchester ** Bryn (ward), an electoral ward in Wigan ** Bryn railway station * Bryn, Shropshire, a location ...
and Gareth W. Griffith in 2020. Prior to being formally classified, ''Myrmecopterula velohortorum'' was referred to as ant cultivar G2 in several studies. It was found in ant nests belonging to the ''Apterostigma'' ''dentigerum'' subclade and was cultivated in hanging 'veiled gardens' where the mycelium formed a thin envelope which surrounded the fungal garden. Gardens are hung under logs or inside cavities within them or rarely found in cavities in the ground. A single hole may exist in the veil serving as the entrance to the nest. It is hypothesized that ''M. velohortorum'' descended from ''M. nudihortorum'' with the two species then taking different evolutionary paths due to co-evolving with ants engaged in varying behaviors. Such as the weaving of mycelial threads to produce the veil which the ants are hypothesized to engage in. ''Myrmecopterula nudihortorum'' was previously referred to as ant cultivar G4 and was found in ant nests belonging to the ''
Apterostigma manni ''Apterostigma'' is a genus of New World ants of the subfamily Myrmicinae. Two species have been described from fossils preserved in Dominican amber, while the others are extant. They are fungus-growing ants, though, unlike the majority of other ...
'' subclade. It is not cultivated in veiled hanging gardens and rather is cultivated in spongelike masses on the bottom of the garden cavity either under logs or in cavities excavated in the ground. The garden is not suspended by a woven veil. This nest building behaviour is more similar to that of lower attine ants which engage in cultivation of Lepiotaceous fungi belonging to the G3 group. Only one species of ''Apterostigma'', ''
Apterostigma auriculatum ''Apterostigma'' is a genus of New World ants of the subfamily Myrmicinae. Two species have been described from fossils preserved in Dominican amber, while the others are extant. They are fungus-growing ants, though, unlike the majority of other ...
'' was documented as cultivating the G3 fungus.


Description

As the 'myrme' prefix in the name ''Myrmecopterula'' would suggest, these species are associated with fungus farming ants found in the
neotropics The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeog ...
of
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. ''Myrmex'' means ants in Greek and
Myrmecology Myrmecology (; from Greek: μύρμηξ, ''myrmex'', "ant" and λόγος, ''logos'', "study") is a branch of entomology focusing on the study of ants. Ants continue to be a model of choice for the study of questions on the evolution of social ...
is the study of ants. The most common fungus known to be cultivated by leafcutter ants is ''
Leucoagaricus gongylophorus ''Leucocoprinus 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 u ...
'' (G1) with little documentation existing about other cultivated species. ''L. gongylophorus'' is cultivated by ''Atta'' and ''
Acromyrmex ''Acromyrmex'' is a genus of New World ants of the subfamily Myrmicinae. This genus is found in South America and parts of Central America, México and the Caribbean Islands, and contains 33 known species. Commonly known as " leafcutter ants" th ...
'' ants, amongst others. ''Myrmecopterula'' fungi however are cultivated by ''
Apterostigma ''Apterostigma'' is a genus of New World ants of the subfamily Myrmicinae. Two species have been described from fossils preserved in Dominican amber, while the others are extant. They are fungus-growing ants, though, unlike the majority of othe ...
'' ants. Unlike ''L. gongylophorus,'' which is no longer capable of spreading via spores, some ''Myrmecopterula'' species may produce sterile or fertile fruiting bodies and may therefore not be as entirely reliant upon the ants. ''M. moniliformis'' produces two distinct forms of mushrooms. The first type resemble irregular strings of beads similar in appearance to some rhizomes produced by plants. These are sterile and lack an active
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in oth ...
to produce spores. The second type are fertile branching coral structures which may grow independently or attached to the sterile bead like structures. These coral like forms are more typical of mushrooms produced by other species in the ''
Pterulaceae The ''Pterulaceae'' are a family (biology), family of fungi in the order Agaricales. According to a 2008 estimate, the family contained 99 species previously distributed among 12 genus, genera. More recent data from molecular phylogenetic reconst ...
'' family. The presence of fertile mushrooms means that ''M. moniliformis'' is capable of reproducing and surviving without the ants. ''M. nudihortorum'' and ''M. velohortorum'' have not been documented as producing fertile fruiting bodies. One explanation may be that mushroom production is actively suppressed by the ants as is seen in ''L. gongylophorus'' which only produces mushrooms when nests are abandoned however active suppression has not been documented in ''Myrmecopterula'' species. This may indicate that ''M. nudihortorum'' and ''M. velohortorum'' have been rendered incapable of producing fertile bodies as a result of prolonged domestication by ants. One hypothesis for the presence of fertile fruiting bodies in ''M. moniliformis'' is that it may have descended from a lineage of ant-domesticated fungi which escaped from cultivation to become free-living fungi. It is still observed as growing on ant nests. The presence of the sterile mushrooms may be a genetic hangover from previous cultivation as this mutation is detrimental to non-domesticated fungi.


Habitat and distribution

''Myrmecopterula'' species are found in the
neotropics The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeog ...
of
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
and are associated with ants of the ''
Apterostigma ''Apterostigma'' is a genus of New World ants of the subfamily Myrmicinae. Two species have been described from fossils preserved in Dominican amber, while the others are extant. They are fungus-growing ants, though, unlike the majority of othe ...
'' genus. Species are usually found growing on top of living or dead ant nests or being cultivated by ants. Some species of ''Myrmecopterula'' grow from soil whilst others appear to grow from wood however closer inspection reveals that rather than using the wood itself as a substrate they are instead found growing from loose debris within cavities in the wood. This substrate is sometimes similar in appearance to that of the fungal gardens of ''
Apterostigma pilosum ''Apterostigma'' is a genus of New World ants of the subfamily Myrmicinae. Two species have been described from fossils preserved in Dominican amber, while the others are extant. They are fungus-growing ants, though, unlike the majority of other ...
'' group ants, which have been documented cultivating ''Myrmecopterula'' species. ''
Apterostigma ''Apterostigma'' is a genus of New World ants of the subfamily Myrmicinae. Two species have been described from fossils preserved in Dominican amber, while the others are extant. They are fungus-growing ants, though, unlike the majority of othe ...
'' species have a distribution which covers South America with some species extending into Central America and Mexico this helps to understand the potential distribution of ''Myrmecopterula'' species. Due to some species of this genus being reliant upon ants and not producing mushrooms, observations are uncommon and would depend on observing the nests themselves. However as ''M. moniliformis'' produces both fertile and infertile mushrooms and can grow without the ants, observations of fruiting bodies can be made which place the distribution around Peru, Brazil, Colombia, Costa Rica and Bolivia. This may help inform the potential distribution of other ''Myrmecopterula'' species.


Etymology

''Myrmecopterula'' derives its name from the Greek ''Myrmex'' for ants and ''Pterula'', the genus it formally belonged to.


Species

At present three ''Myrmecopterula'' species have been named. These are also associated with cultivation by ''
Apterostigma ''Apterostigma'' is a genus of New World ants of the subfamily Myrmicinae. Two species have been described from fossils preserved in Dominican amber, while the others are extant. They are fungus-growing ants, though, unlike the majority of othe ...
'' ants. Four other, unnamed and less well documented species are known.


References

{{Taxonbar, from=Q113003619 Pterulaceae Agaricales genera