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The Zoopagomycotina are a subdivision (''
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 insidians'' with the Kickxellomycotina.
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
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
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, uncertain ...
'') of the fungal division 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 ...
''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 c ...
''. 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 ...
remains in question, so they are sometimes referred to by the informal name zygomycetes.
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 tha ...
parasites
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 ha ...
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- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s, 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.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 c ...
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, ...
. 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 ha ...
(e.g. '' Syncephalis'', '' Piptocephalis'')ygomycetes.org/> that live outside of the host body but produce specialized organs called haustoria 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 (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 ...
. 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 usefu ...
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.comEvolutionary 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 insidians'' with the Kickxellomycotina.
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 unicellu ...
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 chro ...
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 *** FamilyCochlonemataceae
The family Cochlonemataceae is a fungus classified in the order Zoopagales. Fungi found in this family are predominantly parasitic and can be found in any substrate that contains an ample amount of host or prey organisms.
The family contains six ...
Duddington 1974
**** '' Amoebophilus'' (6 spp.) ectoparasite of amoebae
**** '' 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
**** '' Endocochlus'' (4 spp.) endoparasites of amoebae
**** '' Euryancale'' (5 spp. ) endoparasites of nematodes
*** Family Helicocephalidaceae
The Helicocephalidaceae are a family of fungi in the Zoopagales order. The family contain contains 4 genera and 13 species
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as ...
Boedijn 1959
**** '' Brachymyces'' (1 sp.) parasitic on bdelloid rotifers
**** '' 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
The Piptocephalidaceae are a family of fungi in the Zoopagales order. The family contain contains 3 genera, and 70 species
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as ...
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
The ''Sigmoideomycetaceae'' are a family of fungi in the Zoopagales order. The family contain contains three genera, and 4 species
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organ ...
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 Taxonomy (biology), classification and a taxonomic rank of an organism, as well a ...
Drechsler 1938
**** '' Acaulopage'' (27 spp.) haustorial parasites of amoeba
**** '' Cystopage'' (7 spp.) haustorial parasites of amoebae and nematodes
**** '' Lecophagus'' Dick 1990
**** '' Stylopage'' (18 spp.) predaceous on amoebae and nematodes
**** '' Tentaculophagus'' Doweld 2014
**** '' Zoopage'' (11 spp.) haustorial parasite of amoebae and testaceous rhizopods
**** '' Zoophagus'' (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