The Zoopagomycotina are a subdivision (''

incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...

'') of the

fungal 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 th ...

division Division or divider may refer to: Mathematics *Division (mathematics), the inverse of multiplication *Division algorithm, a method for computing the result of mathematical division Military *Division (military), a formation typically consisting ...

Zygomycota Zygomycota, or zygote fungi, is a former division or phylum of the kingdom Fungi. The members are now part of two phyla: the Mucoromycota and Zoopagomycota. Approximately 1060 species are known. They are mostly terrestrial in habitat, living i ...

sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

''. It contains 5 families and 20 genera.ygomycetes.orgurl=http://zygomycetes.org/index.php?id=8 Relationships among and within subphyla of Zygomycota are poorly understood, and their

monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...

remains in question, so they are sometimes referred to by the informal name

zygomycetes Zygomycota, or zygote fungi, is a former phylum, division or phylum of the kingdom Fungi. The members are now part of two Phylum, phyla: the Mucoromycota and Zoopagomycotina, Zoopagomycota. Approximately 1060 species are known. They are mostly t ...

. Zoopagomycotina are microscopic and are typically

obligate {{wiktionary, obligate As an adjective, obligate means "by necessity" (antonym ''facultative'') and is used mainly in biology in phrases such as: * Obligate aerobe, an organism that cannot survive without oxygen * Obligate anaerobe, an organism that ...

parasites Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...

of other zygomycete fungi and of microscopic soil animals such as

nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...

rotifer The rotifers (, from the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals. They were first described by Rev. John H ...

s and

amoebae An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopo ...

.olweb.org/Zygomycota> url=http://www.tolweb.org/Zygomycota Some species are

endoparasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...

that live mostly within the bodies of their hosts and only exit the host when they are producing

spores In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...

. Other species are

ectoparasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...

(e.g. '' Syncephalis'', '' Piptocephalis'')ygomycetes.org/> that live outside of the host body but produce specialized organs called

haustoria In botany and mycology, a haustorium (plural haustoria) is a rootlike structure that grows into or around another structure to absorb water or nutrients. For example, in mistletoe or members of the broomrape family, the structure penetrates t ...

that penetrate inside of the host body to capture host nutrients. Similar haustoria are found in

biotrophic Symbiosis (from Ancient Greek, Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different Organism, biological organisms, be it Mutualism (biolog ...

plant, animal and fungal pathogens in several other major fungal lineages. Like most other zygomycete fungi, the Zoopagomycotina have cell walls containing

chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...

and have

coenocytic A coenocyte () is a multinucleate cell which can result from multiple nuclear divisions without their accompanying cytokinesis, in contrast to a syncytium, which results from cellular aggregation followed by dissolution of the cell membranes insid ...

(nonseptate)

hyphae A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...

. Their vegetative body consists of a simple, branched or unbranched

thallus Thallus (plural: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. Many of these organisms wer ...

. Asexual reproduction is by arthrospores (in '' Helicocephalum''), chlamydospores, uni- or multi-spored sporganiola; sporangiospores of multi-spored formed in simple or branched chains ( merosporangia), usually from a vesicle or stalk. Many produce haustoria. Where observed, the sexual spores ( zygospores) are globose and unornamented. The hyphae used during sexual

outcrossing Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing can be a useful ...

is similar to vegetative hyphae or in some cases may be slightly enlarged.

Etymology

The word Zoopagomycotina comes from the Greek roots ''zoo'' meaning "animal" and ''pag'' meaning "rock" or "ice/frost".ww.oed.comurl=http://www.oed.com

Evolutionary relationships

Although great strides have been made in resolving the evolutionary relationships among many lineages of fungi, it has been challenging to resolve relationships within and among zygomycetes. For example, the uncertain grouping of ''

Zoophagus ''Zoophagus'' is a genus of zygomycete fungi that preys on rotifers and nematodes. It was established in 1911 by Sommerstorff, who originally considered it to be an oomycete. It is common in a variety of freshwater habitats, such as ponds and ...

insidians'' with the

Kickxellomycotina Kickxellomycotina is a fungus grouping. In the subkingdom of Zoopagomyceta . The name was changed from "Harpellomycotina", because "Kickxellomycotina" had an older stem. It came from the genus '' Kickxella'', named after Jean Kickx. Orders in ...

. Resolving a well-supported monophyly of the Zoopagomycotina has been particularly challenging for several main reasons: * most species of Zoopagomycotina are microscopic and challenging to observe * most species of Zoopagomycotina cannot be grown separately from their host organisms in

axenic In biology, axenic (, ) describes the state of a culture in which only a single species, variety, or strain of organism is present and entirely free of all other contaminating organisms. The earliest axenic cultures were of bacteria or unicellul ...

culture, so obtaining pure DNA for molecular studies is challenging * based on

ribosomal DNA Ribosomal DNA (rDNA) is a DNA sequence that codes for ribosomal RNA. These sequences regulate transcription initiation and amplification, and contain both transcribed and non-transcribed spacer segments. In the human genome there are 5 chromos ...

sequences, species of Zoopagomycotina may have undergone accelerated evolution, so grouping may be skewed by

long-branch attraction In phylogenetics, long branch attraction (LBA) is a form of systematic error whereby distantly related lineages are incorrectly inferred to be closely related. LBA arises when the amount of molecular or morphological change accumulated within a lin ...

(LBA) and a high frequency of parallel evolutionary changes

Families and their respective genera

* Class Zoopagomycetes Doweld 2014 ** Order Zoopagales Bessey 1950 ex. Benjamin 1979 oophagales Doweld 2014 *** '' Massartia'' De Wildeman 1897 non Conrad 1926 *** Family Basidiolaceae Doweld 2013 **** '' Basidiolum'' Cienk. 1861 *** Family Cochlonemataceae Duddington 1974 **** '' Amoebophilus'' (6 spp.) ectoparasite of

**** '' Aenigmatomyces'' Castan˜eda & Kendrick 1993 **** '' Aplectosoma'' (1 sp.) parasite of an amoeba **** '' Bdellospora'' (1 sp.) ectoparasite of an amoeba **** '' Cochlonema'' (19 spp.) endoparasites of amoebae and

rhizopods An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of Cell (biology), cell or unicellular organism with the ability to alter its shape, primarily by extending and ret ...

**** '' Endocochlus'' (4 spp.) endoparasites of amoebae **** '' Euryancale'' (5 spp. ) endoparasites of nematodes *** Family Helicocephalidaceae Boedijn 1959 **** '' Brachymyces'' (1 sp.) parasitic on

bdelloid Bdelloidea (Greek ''βδέλλα'', ''bdella'', "leech") is a class of rotifers found in freshwater habitats all over the world. There are over 450 described species of bdelloid rotifers (or 'bdelloids'), distinguished from each other mainly ...

rotifers The rotifers (, from the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals. They were first described by Rev. John H ...

**** '' Helicocephalum'' (5 spp.) parasites of small animals, especially nematodes and nematode eggs **** '' Rhopalomyces'' (8 spp.) parasites of small animals, especially nematodes and nematode eggs **** '' Verrucocephalum'' Degawa 2013 *** Family Piptocephalidaceae Schröter 1886 **** '' Kuzuhaea'' (1 sp.) haustorial parasite of fungi (mostly of Mucorales spp.) **** '' Piptocephalis'' (25 spp.) haustorial parasite of fungi (mostly of Mucorales spp.) **** '' Syncephalis'' (61 spp.) haustorial parasite of fungi (mostly of Mortierellales and Mucorales spp.) *** Family Sigmoideomycetaceae Benny, Benjamin & Kirk 1992 **** '' Reticulocephalis'' (2 spp.) putative haustorial parasite of fungi **** '' Sigmoideomyces'' (2 spp.) putative haustorial parasite of fungi **** '' Sphondylocephalum'' Stalpers 1974 **** '' Thamnocephalis'' (3 spp.) haustorial parasite of fungi *** Family

Zoopagaceae The Zoopagaceae are a family of fungi in the Zoopagales order. The family contain contains 6 genera, and 78 species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodi ...

Drechsler 1938 **** ''

Acaulopage ''Acaulopage'' is a genus in the former Zygomycota that preys on amoeba. Morphology Species of ''Acaulopage'' are characterized by coenocytic hyphae that have patches of yellowish adhesive patches used to capture amoeba. Conidia are borne dir ...

'' (27 spp.) haustorial parasites of amoeba **** '' Cystopage'' (7 spp.) haustorial parasites of amoebae and nematodes **** '' Lecophagus'' Dick 1990 **** ''

Stylopage ''Stylopage'' is a polytypic genus of predacious fungus in the order Zoopagales, within the subphylum Zoopagomycotina. All known species of ''Stylopage'' subsist on various species of amoebae or nematodes by trapping their prey, typically using ...

'' (18 spp.) predaceous on amoebae and nematodes **** '' Tentaculophagus'' Doweld 2014 **** '' Zoopage'' (11 spp.) haustorial parasite of amoebae and testaceous

**** ''

'' (5 spp.) ectoparasites of loricate rotifers and nematodesygomycetes.org/>

References

External links

Zygomycota
at the Tree of Life Web Project
Zygomycetes.org
* * https://www.uniprot.org/taxonomy/451827 * https://web.archive.org/web/20110720003551/http://agclass.canr.msu.edu/mtwdk.exe?w=125844&k=default&s=5&t=2&n=1&l=60 {{Taxonbar, from=Q3131892 Zygomycota