Kickxellales

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Junta Sugiyama - One of the best experts on this subject based on the ideXlab platform.

  • Phylogeny of the Zygomycota based on nuclear ribosomal sequence data
    Mycologia, 2006
    Co-Authors: Merlin M. White, Yuuhiko Tanabe, Kerry O'donnell, Timothy Y. James, Matías J. Cafaro, Junta Sugiyama
    Abstract:

    The Zygomycota is an ecologically heter- ogenous assemblage of nonzoosporic fungi compris- ing two classes, Zygomycetes and Trichomycetes. Phylogenetic analyses have suggested that the phylum is polyphyletic; two of four orders of Trichomycetes are related to the Mesomycetozoa (protists) that diverged near the fungal/animal split. Current circumscription of the Zygomycota includes only orders with representatives that produce zygospores. We present a molecular-based phylogeny including recognized representatives of the Zygomycetes and Trichomycetes with a combined dataset for nuclear rRNA 18S (SSU), 5.8S and 28S (LSU) genes. Tree reconstruction by Bayesian analyses suggests the Zygomycota is paraphyletic. Although 12 clades were identified only some of these correspond to the nine orders of Zygomycota currently recognized. A large superordinal clade, comprising the Dimargaritales, Harpellales, Kickxellales and Zoopagales, grouping together many symbiotic fungi, also is identified in part by a unique septal structure. Although Harpel- lales and Kickxellales are not monophyletic, these lineages are distinct from the Mucorales, Endogo- nales and Mortierellales, which appear more closely related to the Ascomycota + Basidiomycota + Glomeromycota. The final major group, the insect- associated Entomophthorales, appears to be poly- phyletic. In the present analyses Basidiobolus and Neozygites group within Zygomycota but not with the Entomophthorales. Clades are discussed with special reference to traditional classifications, mapping mor- phological characters and ecology, where possible, as a snapshot of our current phylogenetic perspective of the Zygomycota.

  • evolutionary relationships among basal fungi chytridiomycota and zygomycota insights from molecular phylogenetics
    Journal of General and Applied Microbiology, 2005
    Co-Authors: Yuuhiko Tanabe, Makoto M Watanabe, Junta Sugiyama
    Abstract:

    Evolutionary relationships of the two basal fungal phyla Chytridiomycota and Zygomycota are reviewed in light of recent molecular phylogenetic investigation based on rDNA (nSSU, nLSU rDNA), entire mitochondrial genomes, and nuclear protein coding gene sequences (e.g., EF-1α, RPB1). Accumulated molecular evidence strongly suggests that the two basal fungal phyla are not monophyletic. For example, the chytridiomycete order Blastocladiales appears to be closely related to the zygomycete order Entomophthorales. Within the Zygomycota, a monophyletic clade, consisting of the Dimargaritales, Harpellales, and Kickxellales, which is characterized by a shared unique septal ultrastructure, was identified. Moreover, evidence for the exclusion of zygomycete orders Amoebidiales and Eccrinales from the Fungi, and their placement at the Animal-Fungi boundary has been clearly documented. Microsporidia, a group of amitochondriate organisms currently under intensive study, is not supported as derived within the Fungi, but a fungal affinity cannot be ruled out. Taking these molecular phylogenetic studies into account, we proposed a hypothetical evolutionary framework of basal fungi.

  • Molecular phylogeny of Zygomycota based on EF-1α and RPB1 sequences: limitations and utility of alternative markers to rDNA
    Molecular phylogenetics and evolution, 2004
    Co-Authors: Yuuhiko Tanabe, Makoto M Watanabe, Masatoshi Saikawa, Junta Sugiyama
    Abstract:

    Abstract Earlier molecular phylogenetic analyses based on nuclear small subunit ribosomal DNA (nSSU rDNA) suggest that the Zygomycota are polyphyletic within the Chytridiomycota. However, these analyses failed to resolve almost all interordinal relationships among basal fungi (Chytridiomycota and Zygomycota), due to lack of sufficient characters within the nSSU rDNA. To further elucidate the higher-level phylogeny of Zygomycota, we have sequenced partial RPB1 (DNA dependent RNA polymerase II largest subunit) and EF-1α (translation elongation factor 1 alpha) genes from 10 and 3 zygomycete fungi, respectively. Independent molecular phylogenetic analyses were performed based on each sequence by distance and maximum likelihood methods. Although deep phylogenetic relationships among basal fungi still remain poorly resolved using either gene, the RPB1-based phylogeny identified a novel monophyletic clade consisting of the Dimargaritales, Harpellales, and Kickxellales. This result suggests that regularly formed septa (cross walls that divide hyphae into segments) with a lenticular cavity are plesiomorphic for this clade, and indicates the importance of septal pore ultrastructure in zygomycete phylogeny. In addition, a peculiar mucoralean genus Mortierella, which was considered to be distantly related to the other Mucorales based on previous nSSU rDNA analyses, was resolved as the basal most divergence within the Mucorales, consistent with traditional phenotypic-based taxonomy. Although the taxa included in our analysis are restricted, the monophyly of each order suggested by nSSU rDNA phylogeny is supported by the present RPB1-based analysis. These results support the potential use of RPB1 as an alternative marker for fungal phylogenetic studies. Conversely, the overall fungal phylogeny based on EF-1α sequence is poorly resolved. A comparison of numbers of observed substitutions versus inferred substitutions within EF-1α indicates that this gene is much more saturated than RPB1. This result suggests that the EF-1α gene is unsuitable for resolving higher-level phylogenetic relationships within the Fungi.

  • Group I Introns from Zygomycota: Evolutionary Implications for the Fungal IC1 Intron Subgroup
    Journal of Molecular Evolution, 2002
    Co-Authors: Yuuhiko Tanabe, Akira Yokota, Junta Sugiyama
    Abstract:

    The origins of fungal group I introns within nuclear small-subunit (nSSU) rDNA are enigmatic. This is partly because they have never been reported in basal fungal phyla (Zygomycota and Chytridiomycota), which are hypothesized to be ancestral to derived phyla (Ascomycota and Basidiomycota). Here we report group I introns from the nSSU rDNA of two zygomycete fungi, Zoophagus insidians (Zoopagales) and Coemansia mojavensis (Kickxellales). Secondary structure analyses predicted that both introns belong to the IC1 subgroup and that they are distantly related to each other, which is also suggested by different insertion sites. Molecular phylogenetic analyses indicated that the IC1 intron of Z. insidians is closely related to the IC1 intron inserted in the LSU rDNA of the basidiomycete fungus Clavicorona taxophila , which strongly suggests interphylum horizontal transfer. The IC1 intron of C. mojavensis has a low phylogenetic affinity to other fungal IC1 introns inserted into site 943 of nSSU rDNA (relative to E. coli 16S rDNA). It is noteworthy that this intron contains a putative ORF containing a His–Cys box motif in the antisense strand, a hallmark for nuclear-encoded homing endonucleases. Overall, molecular phylogenetic analyses do not support the placement of these two introns in basal fungal IC1 intron lineages. This result leads to the suggestion that fungal IC1 introns might have invaded or been transferred laterally after the divergence of the four major fungal phyla.

  • Molecular Phylogeny of Parasitic Zygomycota (Dimargaritales, Zoopagales) Based on Nuclear Small Subunit Ribosomal DNA Sequences
    Molecular phylogenetics and evolution, 2000
    Co-Authors: Yuuhiko Tanabe, Kerry O'donnell, Masatoshi Saikawa, Junta Sugiyama
    Abstract:

    We analyzed sequence data of the 18S rDNA gene from representatives of nine mycoparasitic or zooparasitic genera to infer the phylogenetic relationships of these fungi within the Zygomycota. Phylogenetic analyses identified a novel monophyletic clade consisting of the Zoopagales, Kickxellales, Spiromyces, and Harpellales. Analyses also identified a monophyletic mycoparasitic–zooparasitic Zoopagales clade in which Syncephalis, Thamnocephalis, and Rhopalomyces form a sister group to a Piptocephalis–Kuzuhaea clade. Although monophyly of the mycoparasitic Dimargaritales received strong bootstrap and decay index support, phylogenetic relationships of this order could not be resolved because of the unusually high rate of base substitutions within the 18S rDNA gene. Overall, the 18S gene tree topology is weak, as reflected by low bootstrap and decay index support for virtually all internal nodes uniting ordinal and superordinal taxa. Nevertheless, the 18S rDNA phylogeny is mostly consistent with traditional phenotypic-based classification schemes of the Fungi.

Yuko Kurihara - One of the best experts on this subject based on the ideXlab platform.

  • Indonesian Kickxellales: two species of Coemansia and Linderina
    Mycoscience, 2008
    Co-Authors: Yuko Kurihara, Nampiah Sukarno, Muhammad Ilyas, Erny Yuniarti, Wibowo Mangunwardoyo, Ju-young Park, Rasti Saraswati, Yantyati Widyastuti, Katsuhiko Ando
    Abstract:

    Four species belonging to Kickxellales (Kickxellomycotina) isolated from soil of Indonesia are described and illustrated. Two new species of Coemansia, C. asiatica and C. javaensis , were discovered in South Sulawesi and West Java, and two known species of Linderina, L. pennispora and L. macrospora , were discovered in East Kalimantan and South Sulawesi, respectively. These four species are newly added to the Indonesian mycobiota. A technique for inducing sporulation of C. javaensis and L. macrospora by adding substances derived from invertebrates such as aphids, nereids, or cladocerans to culture media is described.

  • Pinnaticoemansia, a new genus of Kickxellales, with a revised key to the genera of Kickxellales
    Mycoscience, 2006
    Co-Authors: Yuko Kurihara, Yousuke Degawa
    Abstract:

    Pinnaticoemansia coronantispora gen. sp. nov. (Kickxellaceae, Kickxellales) is described and illustrated. This species is characterized by imparipinnate sporocladia (sporocladia arranged in pairs), sporangiospores with a three-lobed corona, and the germination pattern of sporangiospores with downward hyphal growth and repeated dichotomy. The key to the genera of Kickxellales by Kurihara et al. (2001) is revised based on the observation of the germination pattern of this fungus in comparison with that of Asellariales.

  • Two novel kickxellalean fungi, Mycoëmilia scoparia gen. sp. nov. and Ramicandelaber brevisporus sp. nov.
    Mycological Research, 2004
    Co-Authors: Yuko Kurihara, Yousuke Degawa, Seiji Tokumasu
    Abstract:

    Mycoemilia scoparia gen. sp. nov. is described as a new member of Kickxellales. It is characterized by lageniform sporocladia produced acrogenously in mass and bears wet and fusiform spores on the sporocladia. Ramicandelaber brevisporus sp. nov. is distinguished from the type species of the genus, R. longisporus, by producing much shorter asexual spores, (3-)8(-13) long fertile branches arising from a globose body, and lateral branches.

  • Two novel kickxellalean fungi, Mycoëmilia scoparia gen. sp. nov. and Ramicandelaber brevisporus sp. nov.
    Mycological research, 2004
    Co-Authors: Yuko Kurihara, Yousuke Degawa, Seiji Tokumasu
    Abstract:

    Mycoëmilia scoparia gen. sp. nov. is described as a new member of Kickxellales. It is characterized by lageniform sporocladia produced acrogenously in mass and bears wet and fusiform spores on the sporocladia. Ramicandelaber brevisporus sp. nov. is distinguished from the type species of the genus, R. longisporus, by producing much shorter asexual spores, (3-)8(-13) long fertile branches arising from a globose body, and lateral branches.

  • A new genus Myconymphaea (Kickxellales) with peculiar septal plugs
    Mycological Research, 2001
    Co-Authors: Yuko Kurihara, Yousuke Degawa, Seiji Tokumasu
    Abstract:

    Myconymphaea yatsukahoi gen. sp. nov. is described in the Kickxellaceae (Kickxellales, Zygomycetes). The fungus is characterised by unicellular sporocladia formed on apical vesicles of sporangiophores, conspicuously long sporangiospores and peculiar septal plugs with prominent protuberances. In addition, a key to all known genera of the Kickxellales is provided.

Gerald L. Benny - One of the best experts on this subject based on the ideXlab platform.

  • Preliminary phylogeny of Coemansia (Kickxellales), with descriptions of four new species from Taiwan
    Mycologia, 2017
    Co-Authors: Shu-cheng Chuang, Gerald L. Benny, Nicole Reynolds, Matthew E. Smith, Chiu-yuan Chien, Ju-lin Tsai
    Abstract:

    Four new species of Coemansia from Taiwan are described. Three produce spirally twisted sporangiophores, and these new taxa increase the number of species in the Coemansia spiralis complex from three to six. Each new taxon is morphologically unique. Coemansia biformis, sp. nov., has two different asexual reproductive types on the same thallus; one is straight and the other has a spiral fertile region. Coemansia helicoidea, sp. nov., has stoloniferous sporangiophores with a helicoid fertile region. Coemansia pennisetoides, sp. nov., has a sporangiophore with a fertile region that resembles the inflorescence of the plant genus Pennisetum. Coemansia umbellata, sp. nov., has an umbellate sporangiophore branching pattern and a spirally twisted fertile region on the lowest branches. A dichotomous key was provided to identify the 23 accepted Coemansia species. Phylogenetic analysis based on a combined data set of D1-D2 domains of nuc 28S ribosomal RNA (rDNA) and partial nuc 18S rDNA identifies several independent evolutionary lineages within Coemansia and suggests that Spirodactylon aureum and Kickxella alabastrina may be nested within the genus Coemansia. Sequences of nuc rDNA ITS1-5.8S-ITS2 (internal transcribed spacer [ITS] barcode) are also used to support the description of these new species of Coemansia.

  • Challenges and Future Perspectives in the Systematics of Kickxellomycotina, Mortierellomycotina, Mucoromycotina , and Zoopagomycotina
    Biology of Microfungi, 2016
    Co-Authors: Gerald L. Benny, Eric D. Tretter, Matthew E. Smith, Paul M. Kirk, Merlin M. White
    Abstract:

    This review presents a phylogenetically based classification of four subphyla of the early-diverging fungi: Kickxellomycotina, Mortierellomycotina, Mucoromycotina, and Zoopagomycotina. The Kickxellomycotina contains four orders: Asellariales (symbionts with isopods and Collembola), Dimargaritales (haustorial mycoparasites), Harpellales (symbionts of insect larvae), and Kickxellales (saprobes). Mortierellomycotina contains a single order, Mortierellales, that includes both saprobes and root endophytes. Zoopagomycotina also has one order, Zoopagales, that contains species of obligate animal parasites or mycoparasites. Mucoromycotina has two orders, Endogonales (saprobes and ectomycorrhizal fungi) and Mucorales (primarily saprobes). The Mucorales is by far the most diverse order and includes 14 families and two distinct but unnamed clades. In addition to providing a phylogenetic and taxonomic overview of these subphyla, we provide information on growth and axenic cultivation of these fungi. We also discuss DNA barcoding, environmental sampling, genome sequencing, and phylogenetic analyses as they relate to these subphyla and other early-diverging fungal lineages. Evidence suggests that genome sequencing from a wide array of taxa in these four subphyla combined with innovative environmental sampling approaches is likely to revolutionize our understanding of these fungi and also the fungal tree of life.

  • An eight-gene molecular phylogeny of the Kickxellomycotina, including the first phylogenetic placement of Asellariales
    Mycologia, 2014
    Co-Authors: Eric D. Tretter, Robert W. Lichtwardt, Eric M. Johnson, Gerald L. Benny, Yan Wang, Prasanna Kandel, Stephen J. Novak, James F. Smith, Merlin M. White
    Abstract:

    Kickxellomycotina is a recently described subphylum encompassing four zygomycete orders (Asellariales, Dimargaritales, Harpellales, Kickxellales). These fungi are united by the formation of disciform septal pores containing lenticular plugs. Morphological diversification and life history evolution has made the relationships within and among the four orders difficult to resolve on those grounds alone. Here we infer the phylogeny of the Kickxellomycotina based on an eight-gene supermatrix including both ribosomal rDNA (18S, 28S, 5.8S) and protein sequences (MCM7, TSR1, RPB1, RPB2, β-tubulin). The results of this study demonstrate that Kickxellomycotina is monophyletic and related to members of the Zoopagomycotina. Eight unique clades are distinguished in the Kickxellomycotina, including the four defined orders (Asellariales, Dimargaritales, Harpellales, Kickxellales) as well as four genera previously placed within two of these orders (Barbatospora, Orphella, Ramicandelaber, Spiromyces). Dimargaritales and Ramicandelaber are the earliest diverging members of the subphylum, although the relationship between these taxa remains uncertain. The remaining six clades form a monophyletic group, with Barbatospora diverging first. The next split divides the remaining members of the subphylum into two subclades: (i) Asellariales and Harpellales and (ii) Kickxellales, Orphella and Spiromyces. Estimation of ancestral states for four potentially informative morphological and ecological characters reveals that arthropod endosymbiosis might have been an important factor in the early evolution of the Kickxellomycotina.

  • Phylogenetic relationships among the Harpellales and Kickxellales
    Mycologia, 1998
    Co-Authors: Kerry O'donnell, Elizabeth Cigelnik, Gerald L. Benny
    Abstract:

    DNA sequences of the nuclear encoded small subunit (SSU) 18S ribosomal RNA gene and characters from a morphological/physiological dataset were analyzed by maximum parsimony to investigate phylogene...

Seiji Tokumasu - One of the best experts on this subject based on the ideXlab platform.

Yuuhiko Tanabe - One of the best experts on this subject based on the ideXlab platform.

  • Phylogeny of the Zygomycota based on nuclear ribosomal sequence data
    Mycologia, 2006
    Co-Authors: Merlin M. White, Yuuhiko Tanabe, Kerry O'donnell, Timothy Y. James, Matías J. Cafaro, Junta Sugiyama
    Abstract:

    The Zygomycota is an ecologically heter- ogenous assemblage of nonzoosporic fungi compris- ing two classes, Zygomycetes and Trichomycetes. Phylogenetic analyses have suggested that the phylum is polyphyletic; two of four orders of Trichomycetes are related to the Mesomycetozoa (protists) that diverged near the fungal/animal split. Current circumscription of the Zygomycota includes only orders with representatives that produce zygospores. We present a molecular-based phylogeny including recognized representatives of the Zygomycetes and Trichomycetes with a combined dataset for nuclear rRNA 18S (SSU), 5.8S and 28S (LSU) genes. Tree reconstruction by Bayesian analyses suggests the Zygomycota is paraphyletic. Although 12 clades were identified only some of these correspond to the nine orders of Zygomycota currently recognized. A large superordinal clade, comprising the Dimargaritales, Harpellales, Kickxellales and Zoopagales, grouping together many symbiotic fungi, also is identified in part by a unique septal structure. Although Harpel- lales and Kickxellales are not monophyletic, these lineages are distinct from the Mucorales, Endogo- nales and Mortierellales, which appear more closely related to the Ascomycota + Basidiomycota + Glomeromycota. The final major group, the insect- associated Entomophthorales, appears to be poly- phyletic. In the present analyses Basidiobolus and Neozygites group within Zygomycota but not with the Entomophthorales. Clades are discussed with special reference to traditional classifications, mapping mor- phological characters and ecology, where possible, as a snapshot of our current phylogenetic perspective of the Zygomycota.

  • evolutionary relationships among basal fungi chytridiomycota and zygomycota insights from molecular phylogenetics
    Journal of General and Applied Microbiology, 2005
    Co-Authors: Yuuhiko Tanabe, Makoto M Watanabe, Junta Sugiyama
    Abstract:

    Evolutionary relationships of the two basal fungal phyla Chytridiomycota and Zygomycota are reviewed in light of recent molecular phylogenetic investigation based on rDNA (nSSU, nLSU rDNA), entire mitochondrial genomes, and nuclear protein coding gene sequences (e.g., EF-1α, RPB1). Accumulated molecular evidence strongly suggests that the two basal fungal phyla are not monophyletic. For example, the chytridiomycete order Blastocladiales appears to be closely related to the zygomycete order Entomophthorales. Within the Zygomycota, a monophyletic clade, consisting of the Dimargaritales, Harpellales, and Kickxellales, which is characterized by a shared unique septal ultrastructure, was identified. Moreover, evidence for the exclusion of zygomycete orders Amoebidiales and Eccrinales from the Fungi, and their placement at the Animal-Fungi boundary has been clearly documented. Microsporidia, a group of amitochondriate organisms currently under intensive study, is not supported as derived within the Fungi, but a fungal affinity cannot be ruled out. Taking these molecular phylogenetic studies into account, we proposed a hypothetical evolutionary framework of basal fungi.

  • Molecular phylogeny of Zygomycota based on EF-1α and RPB1 sequences: limitations and utility of alternative markers to rDNA
    Molecular phylogenetics and evolution, 2004
    Co-Authors: Yuuhiko Tanabe, Makoto M Watanabe, Masatoshi Saikawa, Junta Sugiyama
    Abstract:

    Abstract Earlier molecular phylogenetic analyses based on nuclear small subunit ribosomal DNA (nSSU rDNA) suggest that the Zygomycota are polyphyletic within the Chytridiomycota. However, these analyses failed to resolve almost all interordinal relationships among basal fungi (Chytridiomycota and Zygomycota), due to lack of sufficient characters within the nSSU rDNA. To further elucidate the higher-level phylogeny of Zygomycota, we have sequenced partial RPB1 (DNA dependent RNA polymerase II largest subunit) and EF-1α (translation elongation factor 1 alpha) genes from 10 and 3 zygomycete fungi, respectively. Independent molecular phylogenetic analyses were performed based on each sequence by distance and maximum likelihood methods. Although deep phylogenetic relationships among basal fungi still remain poorly resolved using either gene, the RPB1-based phylogeny identified a novel monophyletic clade consisting of the Dimargaritales, Harpellales, and Kickxellales. This result suggests that regularly formed septa (cross walls that divide hyphae into segments) with a lenticular cavity are plesiomorphic for this clade, and indicates the importance of septal pore ultrastructure in zygomycete phylogeny. In addition, a peculiar mucoralean genus Mortierella, which was considered to be distantly related to the other Mucorales based on previous nSSU rDNA analyses, was resolved as the basal most divergence within the Mucorales, consistent with traditional phenotypic-based taxonomy. Although the taxa included in our analysis are restricted, the monophyly of each order suggested by nSSU rDNA phylogeny is supported by the present RPB1-based analysis. These results support the potential use of RPB1 as an alternative marker for fungal phylogenetic studies. Conversely, the overall fungal phylogeny based on EF-1α sequence is poorly resolved. A comparison of numbers of observed substitutions versus inferred substitutions within EF-1α indicates that this gene is much more saturated than RPB1. This result suggests that the EF-1α gene is unsuitable for resolving higher-level phylogenetic relationships within the Fungi.

  • Group I Introns from Zygomycota: Evolutionary Implications for the Fungal IC1 Intron Subgroup
    Journal of Molecular Evolution, 2002
    Co-Authors: Yuuhiko Tanabe, Akira Yokota, Junta Sugiyama
    Abstract:

    The origins of fungal group I introns within nuclear small-subunit (nSSU) rDNA are enigmatic. This is partly because they have never been reported in basal fungal phyla (Zygomycota and Chytridiomycota), which are hypothesized to be ancestral to derived phyla (Ascomycota and Basidiomycota). Here we report group I introns from the nSSU rDNA of two zygomycete fungi, Zoophagus insidians (Zoopagales) and Coemansia mojavensis (Kickxellales). Secondary structure analyses predicted that both introns belong to the IC1 subgroup and that they are distantly related to each other, which is also suggested by different insertion sites. Molecular phylogenetic analyses indicated that the IC1 intron of Z. insidians is closely related to the IC1 intron inserted in the LSU rDNA of the basidiomycete fungus Clavicorona taxophila , which strongly suggests interphylum horizontal transfer. The IC1 intron of C. mojavensis has a low phylogenetic affinity to other fungal IC1 introns inserted into site 943 of nSSU rDNA (relative to E. coli 16S rDNA). It is noteworthy that this intron contains a putative ORF containing a His–Cys box motif in the antisense strand, a hallmark for nuclear-encoded homing endonucleases. Overall, molecular phylogenetic analyses do not support the placement of these two introns in basal fungal IC1 intron lineages. This result leads to the suggestion that fungal IC1 introns might have invaded or been transferred laterally after the divergence of the four major fungal phyla.

  • Molecular Phylogeny of Parasitic Zygomycota (Dimargaritales, Zoopagales) Based on Nuclear Small Subunit Ribosomal DNA Sequences
    Molecular phylogenetics and evolution, 2000
    Co-Authors: Yuuhiko Tanabe, Kerry O'donnell, Masatoshi Saikawa, Junta Sugiyama
    Abstract:

    We analyzed sequence data of the 18S rDNA gene from representatives of nine mycoparasitic or zooparasitic genera to infer the phylogenetic relationships of these fungi within the Zygomycota. Phylogenetic analyses identified a novel monophyletic clade consisting of the Zoopagales, Kickxellales, Spiromyces, and Harpellales. Analyses also identified a monophyletic mycoparasitic–zooparasitic Zoopagales clade in which Syncephalis, Thamnocephalis, and Rhopalomyces form a sister group to a Piptocephalis–Kuzuhaea clade. Although monophyly of the mycoparasitic Dimargaritales received strong bootstrap and decay index support, phylogenetic relationships of this order could not be resolved because of the unusually high rate of base substitutions within the 18S rDNA gene. Overall, the 18S gene tree topology is weak, as reflected by low bootstrap and decay index support for virtually all internal nodes uniting ordinal and superordinal taxa. Nevertheless, the 18S rDNA phylogeny is mostly consistent with traditional phenotypic-based classification schemes of the Fungi.