Ubiquitin in Fungi
Abstract
Ubiquitin is a family member of structural proteins conserved to help in regulating numerous processes within the eukaryotic cells. Such cells contain organelles, nucleus, and enclosed through a plasma membrane. Ubiquitin and other family members perform their functions by a covalent add-on to other cellular proteins. Such attachments alter the stability, activity, or localization of the protein targeted. Such organisms that contain eukaryotic cells include fungi, animals, plants, and protozoa. The ubiquitin amino acid controls each significant movement of proteins within the cell. In the process, they assist in the synthesis of new proteins and destroying the defective ones. The research presents recent literature on mini-article reviews on groups of fungi that include Ascomycota, Cryptomycota, Neocallimastigomycota, Chytridiomycota, Zoopagomycota, Mucoromycota, and Basidiomycota. The mini-article reviews provide information on each member of the Ubiquitin family stated. In this case, the article reviews mainly the ubiquitin in fungi and other groups.
Keywords: Ubiquitin, fungi, proteins and amino acid
Contents
Ascomycota 12
List of Abbreviations
DNA: Deoxyribonucleic acid
E1-Enzyme 1
E2-Enzyme 2
E3- enzyme 3
LUBAC: Linear Ubiquitin chain assembly complex
Introduction
Ubiquitin involves an undersized 76-amino acid, a regulatory protein discovered in 1975. All eukaryotic cells contain ubiquitin that remains in charge of all movements within the cell. The regulatory protein takes part in the synthesizing of new protein as it destroys those not functioning. The ubiquitin found within the eukaryotic cells has a similar amino acid sequence. Evolution has not altered the ubiquitin sequence in the cells. Membranes separate the specialized functions in the cell. Ubiquitin attaches itself to proteins and tags them for disposal, a process known as ubiquitination (Swatek & Komander, 2016). After tagging the proteins, they move to the proteasomes for destruction. Additionally, the ubiquitin detaches itself from the defective protein before entering the proteasomes to enable its reusability. Ubiquitin plays a significant role in the therapy for treating cancer. Doctors utilize ubiquitin in manipulating the protein in cells with cancer-causing the death cells containing cancer. Various studies on ubiquitin have resulted in the formulation of three proteasomes inhibitors that treat individuals with multiple myeloma a type of blood cancer (Wijayawardene, Pawłowska, Letcher, Kirk, Humber, Schüßler & Gęsiorska, 2018). The inhibitors include ixazomib, carfilzomib, and bortezomib inhibitors. The other fungi groups besides ubiquitin include Ascomycota, Cryptomycota, Neocallimastigomycota, Chytridiomycota, Zoopagomycota, Mucoromycota, and Basidiomycota. The research seeks to present mini-article reviews on these groups of fungi and their relation to ubiquitin in fungi.
Ubiquitin
Kirby N. Swatek and David Komander on the issue of Ubiquitin modifications
The review provides overviews on different ubiquitin modifications found within cells highlighting current advancement on ubiquitin chain biology. The review discusses recent results in ubiquitin acetylation and phosphorylation by concentrating on Ser65-phosphorylation and its function in mitophagy and activation of parkin.
The discovery that ubiquitin-protein could modify histones through Lys-linked isopeptide illustrated a new beginning in post-translational signaling. Prior to such discovery, scientists already knew that acetylation and phosphorylation modifications regulated functions of the protein. The 1980s witnessed increased significance and prevalence of protein-based modifications when they connected the ATP-reliability ubiquitination of substrates to their degradation through the 26S proteasomes. Research indicates that over 1000 proteins regulate the process of ubiquitination on human cells (Swatek & Komander, 2016). Ubiquitin attaches itself to substrates through a three-step enzymatic cascade that includes E1 ubiquitin activating, E3 ubiquitin ligating enzymes and E2 ubiquitin conjugating as shown in the figures below. Most proteins experience the ubiquitination process during their cellular lifetime. The process of ubiquitination begins through a single attachment of a ubiquitin molecule in a substrate Lys residue. Figure 1 (Swatek & Komander, 2016).
The figure above shows the modification sites of ubiquitin. Structure A shows the eight sites of the ubiquitination process. Structure B shows six out of seven Lys residues on ubiquitin. Structure C shows identified phosphorylation sites of ubiquitin. The red spheres illustrate phosphorylatable hydroxyl groups on Ser/Thr and Tyr residues.
A Conceptual Overview
Ubiquitin involves a 76-amino acid protein that contains numerous sites for additional post-translational modifications. The seven Lys residues make up the vital features of the ubiquitin. One could ubiquitinate the residues to bring about isopeptide-linked chains of ubiquitin. Proteomic research indicates all linkages available co-exist in cells. Lys48-linked form the most dominant linkage type in cells that target proteins to the proteasomes for degradation (Swatek & Komander, 2016). Recent characterization of the remaining atypical ubiquitin modification has resulted in the discovery of the existence of linkage-specific enzymes and the proteins that assemble hydrolyze and recognize every ubiquitin chain type. Recently, new layers of ubiquitin code have emerged hence enabling the establishment that ubiquitin does not only ubiquitinate but also remain modified through other modifications. Minor chemical post-translational modifications like acetylation and phosphorylation modify ubiquitin. Research indicates that 6 out of 7 ubiquitin Lys residues easily acetylate through modification.
Insights into Ubiquitin Signaling Chain
Current tools and methodologies have given new insights on ubiquitin signaling. Additionally, much progress has taken place on enzymatic machinery of the ubiquitin system and functions of variously linked polyubiquitin signals. Studies on met1-linked linear ubiquitin chains demonstrate how various studies of a specific chain type could advance biology. A plethora of genetic models has established significant roles of linear Ubiquitin chain assembly complex (LUBAC) in immunity and inflammation (Swatek & Komander, 2016). The author indicates that Met1-linked ubiquitin chains serve as vital positive regulators of necrosis factor signaling. Characterization and identification of the cellular machinery that binds assemble and hydrolyze Met1-linked polyubiquitin have promoted the progress made in such studies.
New roles of Lys6-linked polyubiquitin
The author indicates that recent studies have not established clear physiological roles of Lys6-linked chains. In this case, the chains remain enriched upon proteasome inhibition demonstrating non-degradative roles. Studies indicate that Lys6 and Lys33-linkages upregulate upon UV genotoxic stress. Consequently, the author indicates that recently no tools for particular detection of Lys6-linked chains hence bring about the need for further research on the consequences and localization of such modification after the damage of the DNA (Swatek & Komander, 2016). Other studies have linked Lys6-linked chains to the homeostasis of the mitochondrial. Mitophagy needs ubiquitination of the defective mitochondria-mediated largely by the E3 ligase parkin. The utilization of ubiquitin replacement strategy indicates that Lys6 and Lys63 mutations experience delayed mitophagy, demonstrating that the linkage kinds invoke particular downstream processes. Other processes like xenophagy demonstrate the Lys6-linked chains because parkin restricts the intracellular mycobacteria. Besides, studies indicate that E3 ligase bacteria effector, NleL, assembles the Lys6-linkages too. Therefore, a bacterium has the ability to exploit the whole of the ubiquitin code despite the unclearness of the pathophysiological function of the Lys6-linkage ubiquitin. The authors have not indicated any form or kind of conflict of interest in the study of Ubiquitin modifications.
Masato Akutsu, Ivan Dikic, and Anja Bremm, on ubiquitin chain diversity
The authors have indicated that ubiquitin is one of the proteins that play a vital role in the modulation protein function. Additionally, the authors illustrate that ubiquitin deregulation results in the formulation of numerous diseases for humans (Akutsu, Dikic & Bremm, 2016). For this reason, ubiquitin could create different heterotypic and homopolymers on protein substrate resulting in distinct cellular responses. Researchers have indicated that ubiquitin linkage serves in the assembling and propagation of particular signals in vivo. Besides, phosphorylated ubiquitin molecules and ubiquitin chains have acquired much attention in recent times. The authors have illustrated that ubiquitin covalently attaches itself by C-terminus to the protein substrate through the sequential action of various enzymes E1, E2, and E3. The study indicates that multiple or single lysine (K) residues could modify proteins.
Neocallimastigomycota
Wang, Liu, and Groenewald; on the phylum Neocallimastigomycota
The author focuses on the morphology of the Neocallimastigomycota group of fungi. Additionally, the author discusses the molecular phylogeny of Neocallimastigomycota fungi. The Neocallimastigomycota fungi group, according to the authors, contains eight genera of anaerobic fungi. The insufficient sequence data of these genera makes the evolutionary relationships of these genera uncertain (Wang, Liu & Groenewald, 2017). The author indicates that anaerobic fungi serve as the key contributor to plant fiber degradation within the hindgut and rumen of larger herbivorous animals. The Neocallimastigomycota fungi not only serve a huge significance in animal nutrition and rumen function but they also a huge potential to improve lignocelluloses into bio-energy products. Previously researchers described this group of fungi as chytridiomycetous fungi because of their life cycle attributes and biochemistry. However, current scholars classified them under a new family neocallimastigaceae under the order chytridiomycetous because of the similarities of zoospore ultra-structure of some members in the same order.
The Relationships on the Genera
Present studies show that anaeromyces as more distant from the other three genera, while those of others remain unresolved. Additionally, studies indicate that genus promises remain divergent. The new genera’s morphology fits with the genus piromyces but based on genetic, and it seems closer to genus anaeromyces (Wang, Liu & Groenewald, 2017). Consequently, the evolutionary relationships between various genera need further discussion. Various studies have challenged the phylogenic of the Neocallimastigomycota fungi not because of their variable nature of the ITS region, but also due to said species of fungi have no sequence information on their specimen (Wang, Liu & Groenewald, 2017). Furthermore, studies indicate that genetic identification of a good number of Neocallimastigomycota fungi clones remains doubtful. The author also demonstrates that despite huge progress on in both phylogenetic and morphological studies of Neocallimastigomycota fungi, it remains clear that taxonomic backbone that agrees with the botanical name of Nomenclature remain needed urgently to enhance a better comprehension the evolution of such relationships, functions of the organisms in Neocallimastigomycota fungi group and ecology.
Morphology
Morphological and phylogenetic investigation indicates the presence of Neocallimastigomycota species and an orpinomyces species. The Neocallimastigomycota group of fungi’s strict temperature and anaerobic nature needs to make the group more challenging to retain the fungi healthy and viable for further studies (Wang, Liu & Groenewald, 2017). Such a challenge makes it difficult to preserve the pure cultures of anaerobic fungi. Such challenges make it possible to lose the cultures of the species in this group of fungi. The current studies also indicate a broad variation in morphology hence challenging the taxonomic study of Neocallimastigomycota fungi. The author presents the most recent molecular knowledge of Neocallimastigomycota fungi. The fungi appear as shown in figure 2 below
Figure 2(Wang, Liu & Groenewald, 2017).
The figure shows the morphology of anaerobic fungi from yak; a thallus, b zoospore, Orpinomyces; c thallus, d zoospore
The author indicates that the study lacks complete credibility since the researchers lacked LSU sequences of the isolated isolates that could have helped in establishing their position within the LSU phylogeny.
Ascomycota
Egbuta, Mwanza, and Babalola, on Ubiquity of Ascomycetes filamentous fungi and their medical and economic significance
The study seeks to review the current literature on the Ascomycota fungi group. Additionally, the study presents the ubiquity of Ascomycota fungi in relation to their medical and economic significance. The key groups on the Ascomycota group of fungi include taphrinomycotina that lack ascomata and recognized recently through molecular evidence provided by Sugiyama and Nishida in 1994 (Egbuta, Mwanza & Babalola, 2016). The basal nature of the Ascomycota fungi remains illustrated from the molecular phylogenetic evidence, morphology, and simple life. The other group involves the Saccharomycotina that lacks ascomata and the unitunicate asci. The last category involves the pezizomycotina that appears when the sexual stage establishes itself. Additionally, this category has a more complex morphology that includes lichen-forming fungi.
Filamentous is one of the fungi among the Ascomycota fungi group found in the soil, water, and air, according to the authors’ research. The group contains numerous organisms from various classes based on the sub-phylum pezizomycotina (Egbuta, Mwanza & Babalola, 2016). The distribution of the filamentous fungi serves numerous purposes. Recent studies indicate that Ascomycota fungi contain cell walls that have chitin that separates them from place cell walls that have bacterial and cellulose cell walls. An Ascomycota fungus occurs naturally in the environment, and they remain unseen because of their cryptic lifestyles on certain substrates they inhabit. The filamentous fungi exist naturally in the different ecosystems, and the environment remains as the only one known fungi.
Economic and medical significance
The Fusarium fungi under the filamentous group mainly cause a broad range of diseases and infections in animals and humans. Furthermore, they help in the production of conidia in a penicillus structure posing a huge economic significance between aspergillus and penicillium (Egbuta, Mwanza & Babalola, 2016). However, various studies indicate that the Ascomycota fungi group serves as a raw material in the agricultural and manufacturing sectors globally. They form a source of raw materials for chemicals, food, cosmetic and pharmaceutical industries. Current studies indicate that filamentous fungi from the Ascomycota group serve as bioremediation agents that degrade the contents of highly contaminated soils chemically hence decreasing toxicity of the soil.
Besides, the Ascomycota group might help in the production of bio-fuel. The same has negative implications like decreasing the food crop’s nutritional value as well as the economic consequences. The manufacturing industry utilizes the Ascomycota fungi to manufacture paper utilized for writing and printing. The food industry utilizes fungi during the fermentation process (Egbuta, Mwanza & Babalola, 2016). Further studies indicate that the filamentous fungi under the phylum Ascomycota could have either a positive or negative effect on animals and humans. In this case, the group of fungi produces various metabolites that contain various inhibitory consequences in the metabolic pathways. The fungi inhibit microbial growth by producing pyrrocidines inform of effective antibiotics against gram-positive bacteria. The authors have clearly stated that the study has no conflict of interest.
Basidiomycota
Dewi, Aryantha and Kandar; on the diversity of Basidiomycota Fungi as a source of Nutraceutical
The author seeks to acquire Basidiomycota fungi isolates that have the ability as a Nutraceutical source. Consequently, the study seeks to establish the diversity of the Basidiomycota group of fungi. According to the authors, recent studies indicate that data concerning fungi diversity and its use remain nearly 712,000 species (Dewi, Aryantha & Kandar, 2018). Microbial technology demonstrates the diversification of agriculture by large-scale recycling of the agro-wastes. Studies have also indicated that button mushrooms, shiitake mushrooms, and oyster mushrooms serve as food and have bioactive elements that serve the source of Nutraceutical. The cultivation of the fungi allows for broad diversity, but few remain cultivated successfully. Most countries around the world have not started the cultivation of Basidiomycota fungi.
Division of Basidiomycota traits
The authors indicate that there exist over 70,000 Basidiomycota fungi of the total 1,500,000 species globally. Countries such as Indonesia have increased diversity of animals and plants as well as mushrooms because the tropical temperatures and humid surroundings support fungi growth (Dewi, Aryantha & Kandar, 2018). The author illustrates that the type of fungi determines the structure of the fungi. The Basidiomycota fungi have a single cell. Humans could eat some Basidiomycota fungi, but some are poisonous such as those from the Aminata genus. Some of the Basidiomycota fungi would poison crops, whereas others from the genus Auricularia, Pleurotus, and Volvariella serve as food to humans.
Other scholars have established that the color of the mushroom helps in identifying the poisonous one and the one suitable for consumption. The main characteristic of the Basidiomycota fungi is that they lack chlorophyll. Such a trait forms its way of life as a parasite. The body contains branching threads known as hyphae. The author indicates that a group of hyphae forms a mycelium that reproduces both sexually and asexually. The mycelium puts artificial threads on the fruiting body of the fungi (Dewi, Aryantha & Kandar, 2018). Additionally, the hyphae entail threadlike structures made of pipe-shaped walls that cover the cytoplasm and plasma membrane of hyphae. The cytoplasm has eukaryotic organelles. The transverse septa or walls have a huge pore sufficient to pass through the mitochondria, ribosome, and the nucleus of the cell that moves from one cell to another cell.
The author calls those not septic senositic hyphae. The authors indicate that numerous divisions of the nucleus cell that not tracked by the cytoplasmic division cause the senocytic hyphae structure. The hyphae also have a clamp connection that demonstrates a trait of fungi belonging to the Basidiomycota class. Furthermore, the authors indicate that Basidiomycota mushrooms form part of the most nutritious food free of cholesterol. Other research studies report that a mushroom has 14-35% protein, 72% unsaturated fat, and 100kj/100g calories (Dewi, Aryantha & Kandar, 2018). The Basidiomycota fungi contain nine key amino acids in the body. Lastly, the Basidiomycota fungi serve as an antioxidant, immunomodulator, anti-inflammatory, and anti-cancer. The authors have indicated there is no conflict of interest
Mucoromycota
“Ying Chang and Gregory Michael Bonito: On phylogenomics of endogonaceae and mycorrhizas evolution within Mucoromycota fungi.
The authors indicate that mucoromycotina contains densosporaceae and endogonaceae that form the only identified non-dikarya order among the ectomycorrhozal members. Such fungi also form a mycorrhizal-like relation with other non-spermatophyte plants. Mycorrhizal fungi in the Mucoromycota category give a host plant with numerous nutrients like calcium and manganese. The Mucoromycota fungi receive carbon in the form of glucose. Therefore, Mucoromycota fungi help in the cycling of nutrients in terrestrial ecosystems (Chang, Desirò, Na, Sandor, Lipzen, Clum & Smith, 2019). The author also indicates that arbuscular mycorrhizas of mucorimycota and ectomycorrhizas formulated by the family members of Basidiomycota and Ascomycota from the main groups of mycorrhizal symbioses among the Mucoromycota fungi. The speedy sequencing of Mucoromycota genomes over the last few years has extremely expanded the knowledge on the diversity of the fungi tree life, increasing the comprehension of fungal biology. Research indicates that eukaryotes related to fungal sporocarps might exist as symbionts, as predictable members of the sporocarp microbiome or as environmental contaminants. Besides, bioinformatics acquiring of metagenomic information is the key obstacle in acquiring access to the fungal genome from a sporocarp tissue.
Zoopagomycota
William J. Davis and Kevin R. Amses, on Genome-scale phylogenetics, shows monophyletic Zoopagomycota fungi
Previous studies have shown that genome-scale phylogenetic examination of fungi has sampled fewer taxa from Zoopagomycota fungi like zoopagales that contains numerous parasitic genera. Besides, such fungi never grow in the pure culture environment. The authors illustrate that Zoopagomycota fungi form an evolutionary pedigree in the opishthokonta with remarkable ecological and morphological diversity (Davis, Amses, Benny, Carter-House, Chang, Grigoriev & James, 2019). Such fungi involve single-celled organisms and multi-cellular organisms that have differentiated tissues. The Zoopagomycota fungi form the most suitable group of fungi to study on the evolution of interactions and complex traits like a predacious lifestyle. The authors indicate that the nodes within fungi remain under-sampled hence limiting the capability to rebuild ancestral traits and establish homologies of the lifestyles within the Zoopagomycota fungi specifically.
Zygomycetes represent one of the earliest fungi forms that transformed from Chytridiomycota and Blastocladiomycota to non-flagellated taxa with an initial aerial dispersal and terrestrial habit. Zoopagomycota fungi relate to heterotrophic eukaryotes. The authors indicate that Zoopagomycota has three subphyla that include zoopagomycotina, kickxellomycotina, and entomophthoramycotina. Various scholars have indicated that most people have not concentrated on both zoopagomycotina and Zoopagomycota (Davis, Amses, Benny, Carter-House, Chang, Grigoriev & James, 2019). On the other hand, many scholars have studied zoopagales fungi due to their diversity of predacious fungal genera. Studies confirm that zoopagales is one of the monophyletic lineages. Nevertheless, the relationship of zoopagales with other subphyla and orders has remained unresolved. Furthermore, present studies indicate that genera zoopagales needs reclassification. The authors indicate that there exists no conflict of interest.
Cryptomycota
Lene Lange, Kristian Barret, Bo Pilgaard, Frank Gleason, Adrian Tsang, on enzymes of early-diverging, zoosporic fungi
The authors indicate Cryptomycota forms the oldest diverging lineages in the fungal kingdom. The urge to comprehend the foundation for the devastating chytridiomycosis among amphibians and the function of fungi within the rumen has driven the functional studies of early-diverging lineages of fungi. The willingness of scientists to broaden the fungal genome project has remained significant in the study of the zoosporic fungi. Straightening out the basal lineages of fungal phylogeny forms a prerequisite to comprehending fungi evolution (Lange, Barrett, Pilgaard, Gleason & Tsang, 2019). Early diverging Cryptomycota fungi thrive in numerous habitats like aquatic, terrestrial, aerobic, and anaerobic. Such fungi have also specialized in various lifestyles that include biotrophic, symbiotic, saprotrophic, pathogenic and parasitic. Consequently, the Cryptomycota fungi mobilize organic matters in plants, animals, and algal. The authors have indicated that the early-diverging of Cryptomycota digest many key types of biopolymers. On the other hand, the zoosporic fungi remain inadequate in acting on enzymes on polymer lignin.
Chytridiomycota
Miguel A. Naranjo-Ortiz and Toni Gabaldon, on Fungal evolution: taxonomy, diversity, and phylogeny of the fungi
The author indicates that the Chytridiomycota fungi remain divided into three key classes that include hyaloraphidiomycetes, monoblepharidomycetes, and chytridiomycetes. In this case, the authors indicate that chrytrids provide a zoosporic propagation stage. Additionally, chrytrids cell at times presents various degrees of apical growth. Chytridiomycota fungi are the largest form of fungi among the zoosporic fungi, with approximately 1000 species. Recent studies demonstrate that many lineages have grown to the degree of orders (Naranjo‐Ortiz & Gabaldón, 2019). Monoblepharidomycetes fungi under the Chytridiomycota fungi involve freshwater zoosporic fungi, which could result in mycelia or unicellular growth. In this case, according to the authors, mycelia monoblepharidomycetes create a monophyletic clade in this specific class. This kind of fungi is the only fungi class that forms real hyphae presenting exceptional cytological traits like the presence of centrioles and the lack of Spitzenkorper. The presence of morphologically various gametes remains common among the monoblepharidomycetes hence demonstrating a unique trait among other fungi in the Fungal Kingdom. The authors also indicate that scholars have not intensified the study of hyaloraphidium fungi since it has an unclear lifestyle. In this case, previous studies categorized the fungi as colorless green algae. Nevertheless, molecular phylogenies illustrate affinity when it comes to monoblepharidomycetes. The authors have not indicated any form of conflict of interest in the study.
Conclusion
Ubiquitin involves a structural protein that regulates many processes in the eukaryotic cells. The protein has a nucleus, organelles, and covered by a plasma membrane. The organisms in eukaryotic cells involve animals, fungi, plants, and protozoa. The ubiquitin amino acid pedals every important movement of proteins in a cell. The class of fungi in the fungal kingdom includes but not limited to Ascomycota, Cryptomycota, Neocallimastigomycota, Chytridiomycota, Zoopagomycota, Mucoromycota and Basidiomycota. Various scholars have carried out studies on the Fungal Kingdom, reclassifying many other fungi into new classes and orders. Some fungi have both economic and medicinal value when utilized by humans or plants.
References
Akutsu, M., Dikic, I., & Bremm, A. (2016). Ubiquitin chain diversity at a glance. J Cell Sci, 129(5), 875-880.
Chang, Y., Desirò, A., Na, H., Sandor, L., Lipzen, A., Clum, A., … & Smith, M. E. (2019). Phylogenomics of Endogonaceae and evolution of mycorrhizas within Mucoromycota. New Phytologist, 222(1), 511-525.
Davis, W. J., Amses, K. R., Benny, G. L., Carter-House, D., Chang, Y., Grigoriev, I., … & James, T. Y. (2019). Genome-scale phylogenetics reveals a monophyletic Zoopagales (Zoopagomycota, Fungi). Molecular phylogenetics and evolution, 133, 152-163.
Dewi, M., Aryantha, I. N. P., & Kandar, M. (2018). The Development of Basidiomycota Fungi with The Concept of Integrated Forest Management. Asia Proceedings of Social Sciences, 2(1), 28-32.
Egbuta, M. A., Mwanza, M., & Babalola, O. O. (2016). A review of the ubiquity of ascomycetes filamentous fungi in relation to their economic and medical importance. Advances in Microbiology, 6(14), 1140-1158.
Lange, L., Barrett, K., Pilgaard, B., Gleason, F., & Tsang, A. (2019). Enzymes of early-diverging, zoosporic fungi. Applied microbiology and biotechnology, 103(17), 6885-6902.
Naranjo‐Ortiz, M. A., & Gabaldón, T. (2019). Fungal evolution: diversity, taxonomy and phylogeny of the Fungi. Biological Reviews, 94(6), 2101-2137.
Swatek, K. N., & Komander, D. (2016). Ubiquitin modifications. Cell research, 26(4), 399-422.
Wang, X., Liu, X., & Groenewald, J. Z. (2017). Phylogeny of anaerobic fungi (phylum Neocallimastigomycota), with contributions from yak in China. Antonie Van Leeuwenhoek, 110(1), 87-103.
Wijayawardene, N. N., Pawłowska, J., Letcher, P. M., Kirk, P. M., Humber, R. A., Schüßler, A., … & Gęsiorska, A. (2018). Notes for genera: basal clades of Fungi (including Aphelidiomycota, Basidiobolomycota, Blastocladiomycota, Calcarisporiellomycota, Caulochytriomycota, Chytridiomycota, Entomophthoromycota, Glomeromycota, Kickxellomycota, Monoblepharomycota, Mortierellomycota, Mucoromycota, Neocallimastigomycota, Olpidiomycota, Rozellomycota and Zoopagomycota). Fungal Diversity, 92(1), 43-129.