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href="/report/chapter/hadakaviridae/hadakaviridae/hadakavirus" hreflang="en">Genus: Hadakavirus</a> </li> </ul> </li> <li class="menu-item"> <a href="/report/chapter/hadakaviridae/hadakaviridae/authors" hreflang="en">Authors: Hadakaviridae</a> </li> <li class="menu-item"> <a href="/report/chapter/hadakaviridae/hadakaviridae/citation" hreflang="en">Citation: Hadakaviridae</a> </li> <li class="menu-item"> <a href="/report/chapter/hadakaviridae/hadakaviridae/references" hreflang="en">References: Hadakaviridae</a> </li> <li class="menu-item"> <a href="/report/chapter/hadakaviridae/hadakaviridae/resources" hreflang="en">Resources: Hadakaviridae</a> </li> <li class="menu-item"> <a href="/report/chapter/hadakaviridae/taxonomy/hadakaviridae" hreflang="en">Species List: Hadakaviridae</a> </li> </ul> </div> </div> </div> </div> <div class="col-lg-9 col-md-9 col-12 col-9right"> <div class="_none clearfix block block-layout-builder block-field-blocknodemt-servicefield-mt-srv-body"> <div class="content"> <div class="clearfix text-formatted field field--name-field-mt-srv-body field--type-text-with-summary field--label-hidden field__item"><h2>Family: <em>Hadakaviridae</em></h2> <p><strong>Yukiyo Sato, Massimo Turina, Sotaro Chiba, Ryo Okada, Muhammad F. Bhatti, Ioly Kotta-Loizou, Robert H. A. Coutts, Hideki Kondo, Sead Sabanadzovic and Nobuhiro Suzuki</strong></p> <p>The citation for this ICTV Report chapter is the summary published as Sato <em>et al.,</em> (2023):<br> ICTV Virus Taxonomy Profile: <em>Hadakaviridae </em>2023, Journal of General Virology (2023) 104:001820</p> <p><strong><strong>Corresponding author:</strong></strong> Nobuhiro Suzuki (<a href="mailto:nsuzuki@okayama-u.ac.jp">nsuzuki@okayama-u.ac.jp</a>)<br> <strong><strong>Edited by:</strong></strong> Sead Sabanadzovic and Peter Simmonds<br> <strong><strong>Posted:</strong></strong> November 2022</p> <h2>Summary</h2> <p><em>Hadakaviridae</em> is a family of capsidless positive-sense (+) single-stranded RNA viruses with a genome composed of ten or eleven segments (Table 1.<em>Hadakaviridae</em>). “Hadaka”, meaning “naked” in Japanese and used to name the virus family, describes their unusual capsidless natures. The lack of a capsid means that these viruses cannot be pelleted by conventional ultracentrifugation and their genome and replicative form double-stranded RNAs are susceptible to exogenously added ribonuclease in host tissue homogenates. Currently known hosts are ascomycetous filamentous fungi. Hadakavirids are most closely related to members of the family <em>Polymycoviridae</em> and show phylogenetic affinity to (+) RNA viruses in the phylum <em>Pisuviricota</em> (extended “picornavirus supergroup”).</p> <p><strong>Table 1. <em>Hadakaviridae. </em>Characteristics of members of the family <em>Hadakaviridae </em></strong></p> <table border="1"> <tbody> <tr> <td width="120"> <p><strong>Characteristic</strong></p> </td> <td width="343"> <p><strong>Description</strong></p> </td> </tr> <tr> <td width="120"> <p>Example</p> </td> <td width="343"> <p>hadaka virus 1 (LC519840–LC519850), species <em>Hadakavirus nanga</em> genus <em>Hadakavirus</em></p> </td> </tr> <tr> <td width="120"> <p>Virion</p> </td> <td width="343"> <p>Capsidless (no known virions)</p> </td> </tr> <tr> <td width="120"> <p>Genome</p> </td> <td width="343"> <p>Multi-segmented genome of ten or eleven linear, positive-sense (+) RNA segments of 0.9–2.5 kb, comprising 14–15 kb in total</p> </td> </tr> <tr> <td width="120"> <p>Replication</p> </td> <td width="343"> <p>Double-stranded replicative forms accumulate abundantly in infected fungal hosts. The cellular replication site remains unknown.</p> </td> </tr> <tr> <td width="120"> <p>Translation</p> </td> <td width="343"> <p>From monocistronic non-polyadenylated genomic RNAs</p> </td> </tr> <tr> <td width="120"> <p>Host range</p> </td> <td width="343"> <p>Fungi</p> </td> </tr> <tr> <td width="120"> <p>Taxonomy</p> </td> <td width="343"> <p>Realm <em>Riboviria</em>; kingdom <em>Orthornavirae</em>, phylum <em>Pisuviricota</em>; The family includes one genus <em>Hadakavirus</em> accommodating one species</p> </td> </tr> </tbody> </table> <h2>Virion</h2> <h4>Morphology</h4> <p>Hadakavirids (members of the family <em>Hadakariviridae</em>) have no true virion, and they cannot be purified by conventional virus purification methods (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/34313859" target="ictvref" title=" Khan et al., 2021, A second capsidless hadakavirus strain with 10 positive-sense single-stranded RNA genomic segments from Fusarium nygamai, Arch Virol, 166, 10, 2711-22">Khan <i>et al.,</i> 2021</a>). Their infectious entity remains unknown.</p> <h4>Physicochemical and physical properties</h4> <p>Hadakavirids cannot be pelleted by ultracentrifugation at 100,000 × <em>g</em> for 1.5 h at 4°C, conditions that can normally be used to purify encapsidated viruses independently of their virion size or morphology (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>).</p> <h4>Nucleic acid</h4> <p>The genome of hadaka virus 1 (HadV1) consists of ten or eleven positive-sense (+) RNA segments (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/34313859" target="ictvref" title=" Khan et al., 2021, A second capsidless hadakavirus strain with 10 positive-sense single-stranded RNA genomic segments from Fusarium nygamai, Arch Virol, 166, 10, 2711-22">Khan <i>et al.,</i> 2021</a>). Colletotrichum fructicola RNA virus 1 (CfRV1), an unassigned member of the family <em>Hadakaviridae</em>, has only seven genomic segments (<a href="https://www.ncbi.nlm.nih.gov/pubmed/35435725" target="ictvref" title=" Fu et al., 2022, A novel heptasegmented positive-sense single-stranded RNA virus from the phytopathogenic fungus Colletotrichum fructicola, J Virol, 96, 9, e0031822">Fu <i>et al.,</i> 2022</a>). Double-stranded (ds) RNA replicative forms accumulate abundantly in host fungal mycelia. Both, the hadakavirid genomic (+) RNA and dsRNAs, are susceptible to ribonuclease A (exogenously added) in host tissue homogenates buffered with a neutral phosphate (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/34313859" target="ictvref" title=" Khan et al., 2021, A second capsidless hadakavirus strain with 10 positive-sense single-stranded RNA genomic segments from Fusarium nygamai, Arch Virol, 166, 10, 2711-22">Khan <i>et al.,</i> 2021</a>). How these RNAs are protected <em>in vivo </em>from endogenous host nucleases remains to be determined.</p> <h4>Proteins</h4> <p>No structural proteins have been detected experimentally and/or bioinformatically (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/34313859" target="ictvref" title=" Khan et al., 2021, A second capsidless hadakavirus strain with 10 positive-sense single-stranded RNA genomic segments from Fusarium nygamai, Arch Virol, 166, 10, 2711-22">Khan <i>et al.,</i> 2021</a>). The hypothetical proteins encoded by the hadakavirid genome show no homology to known viral structural proteins.</p> <h4>Lipids</h4> <p>CfRV1 appears to be associated with the formation of giant vesicles (196–559 nm) containing smaller vesicles (25–40 nm) in host fungal cells (<a href="https://www.ncbi.nlm.nih.gov/pubmed/35435725" target="ictvref" title=" Fu et al., 2022, A novel heptasegmented positive-sense single-stranded RNA virus from the phytopathogenic fungus Colletotrichum fructicola, J Virol, 96, 9, e0031822">Fu <i>et al.,</i> 2022</a>). These vesicles appear to be lipid -layered. However, it remains to be determined whether these vesicles contain viral components.</p> <h2>Genome organization and replication</h2> <p>The exemplar virus isolate (HadV1-7n) has an eleven-segmented (+) RNA genome, each containing a single open reading frame (ORF) (Figure 1.<em>Hadakavirida</em><em>e</em>), while hadaka virus 1-1NL (HadV1-1NL) has ten monocistronic segments (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/34313859" target="ictvref" title=" Khan et al., 2021, A second capsidless hadakavirus strain with 10 positive-sense single-stranded RNA genomic segments from Fusarium nygamai, Arch Virol, 166, 10, 2711-22">Khan <i>et al.,</i> 2021</a>). All genome segments start with the conserved three nucleotides 5′-CGUand end with CCA-3′ or CCG-3′. Exceptions to this rule are RNA7 of HadV1-7n and RNA6 of HadV1-1NL, which have the termini 5′-CGU and GGG-3′. Genomic segments are vertically transmitted through asexual spores in an all-or-none fashion. The largest three genomic segments, RNAs1–3, encode proteins homologous to those encoded by dsRNAs1–3 of members of the family <em>Polymycoviridae</em>, namely an RNA-dependent RNA polymerase (RdRP), a hypothetical protein with unknown functions, and a putative methyltransferase (MTR). A unique genomic segment, RNA8, of HadV1-7n encodes a short C<sub>2</sub>H<sub>2</sub>-type zinc finger protein. This genomic segment is missing for HadV1-1NL (<a href="https://www.ncbi.nlm.nih.gov/pubmed/34313859" target="ictvref" title=" Khan et al., 2021, A second capsidless hadakavirus strain with 10 positive-sense single-stranded RNA genomic segments from Fusarium nygamai, Arch Virol, 166, 10, 2711-22">Khan <i>et al.,</i> 2021</a>). Hypothetical proteins encoded by the remaining genomic segments have no significant homology to other known proteins. The other potential member of the <em>Hadakaviridae</em>, CfRV1, has fewer (seven) genomic segments (<a href="https://www.ncbi.nlm.nih.gov/pubmed/35435725" target="ictvref" title=" Fu et al., 2022, A novel heptasegmented positive-sense single-stranded RNA virus from the phytopathogenic fungus Colletotrichum fructicola, J Virol, 96, 9, e0031822">Fu <i>et al.,</i> 2022</a>) with the three largest also encoding proteins homologous to those of polymycovirids. Most CfRV1 genomic segments possess the conserved terminal sequences 5′-CGU and CCN-3′.</p> <table border="&quot1""> <tbody> <tr> <td><img alt="Hadakaviridae genome" data-entity-type="file" data-entity-uuid="d70bd00a-8061-4a99-81c6-2e797ce7f62a" src="/system/files/inline-images/OPSR.Hadaka.Fig1_.v4.png" width="1975" height="1427" loading="lazy"></td> </tr> <tr> <td> <p><strong>Figure 1. </strong><em><strong>Hadakaviridae. </strong></em>Genome organization of hadaka virus 1 isolate 7n (LC519840–50). Open reading frames are indicated by boxes. The three genes homologous between hadakavirids and polymycovirids are coloured pink: P1 (RdRP - RNA-dependent RNA polymerase), P2 (unknown function) and P3 (MTR- methyltransferase). Other colored regions indicate conserved domains or motifs identified previously (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>). The conserved protein domains and motifs were detected from the Conserved Domain Database (<a href="https://www.ncbi.nlm.nih.gov/pubmed/31777944" target="ictvref" title=" Lu et al., 2020, CDD/SPARCLE: the conserved domain database in 2020, Nucleic Acids Res, 48, D1, D265-8">Lu <i>et al.,</i> 2020</a>).</p> </td> </tr> </tbody> </table> <h2>Biology</h2> <p>Known hosts for the hadakavirids are ascomycetous filamentous fungi. HadV1-7n and -1NL were isolated from <em>Fusarium oxysporum</em> and <em>F. nygamai</em>, respectively, in Pakistan (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/34313859" target="ictvref" title=" Khan et al., 2021, A second capsidless hadakavirus strain with 10 positive-sense single-stranded RNA genomic segments from Fusarium nygamai, Arch Virol, 166, 10, 2711-22">Khan <i>et al.,</i> 2021</a>). Neither of these HadV1 isolates have any apparent effect on the host on potato dextrose agar (PDA) media. CfRV1 was discovered from <em>Colletotrichum fructicola</em> in China (<a href="https://www.ncbi.nlm.nih.gov/pubmed/35435725" target="ictvref" title=" Fu et al., 2022, A novel heptasegmented positive-sense single-stranded RNA virus from the phytopathogenic fungus Colletotrichum fructicola, J Virol, 96, 9, e0031822">Fu <i>et al.,</i> 2022</a>). CfRV1 causes a mild growth reduction of the host on PDA medium and an infection delay on harvested pear fruits. No experimental method for infecting the host with extracellular hadakavirids has been developed.</p> <h4>Antigenicity</h4> <p>No antibody against any proteins of hadakavirids has been reported.</p> <h2>Derivation of names</h2> <p><em>Hadakaviridae</em><em><u>:</u></em> derived from <em><u>Hadaka</u></em><em> </em>(裸、はだか), a Japanese word meaning “naked”.</p> <p><em>Hadakavirus nanga</em>: derived from <em><u>nanga</u></em><u> </u>(ننگا), an Urdu word meaning “naked”, Urdu being the official language of Pakistan where HadV1-7n and HadV1-1NL were isolated.</p> <h2>Genus demarcation criteria</h2> <p>No genus demarcation criteria have been defined since the family <em>Hadakaviridae</em> currently consists of one single genus.</p> <h2>Relationships within the family</h2> <p>The family includes a single genus <em>Hadakavirus</em> which accommodates one species, <em>Hadakavirus nanga</em>, with two well-characterized virus isolates HadV1-7n and -1NL (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/34313859" target="ictvref" title=" Khan et al., 2021, A second capsidless hadakavirus strain with 10 positive-sense single-stranded RNA genomic segments from Fusarium nygamai, Arch Virol, 166, 10, 2711-22">Khan <i>et al.,</i> 2021</a>). CfRV1 is phylogenetically closer to HadV1-7n and HadV1-1NL than to polymycovirids (Figure 2.<em>Hadakaviridae</em>) (<a href="https://www.ncbi.nlm.nih.gov/pubmed/35435725" target="ictvref" title=" Fu et al., 2022, A novel heptasegmented positive-sense single-stranded RNA virus from the phytopathogenic fungus Colletotrichum fructicola, J Virol, 96, 9, e0031822">Fu <i>et al.,</i> 2022</a>).</p> <table border="&quot1""> <tbody> <tr> <td><img alt="Hadakaviridae phylogeny" data-entity-type="file" data-entity-uuid="e450d51f-83a4-4acb-a2d0-ebebf45218df" src="/system/files/inline-images/OPSR.Hadaka.Fig2_.v1.png" width="4115" height="1185" loading="lazy"></td> </tr> <tr> <td> <p><strong>Figure 2.</strong><em><strong>Hadakaviridae. </strong></em>Phylogenetic tree of hadakavirids and related viruses. Members of the same genus have tips labelled with the magenta (<em>Hadakvirus</em>) or cyan (<em>Polymycovirus</em> – used as an outgroup); an unassigned virus as an open circle. The maximum likelihood tree was generated based on the amino acid sequence alignment of RdRP. Multiple sequence alignment was performed by the L-INS-i method in MAFFT v7.490 (<a href="https://www.ncbi.nlm.nih.gov/pubmed/23329690" target="ictvref" title=" Katoh and Standley 2013, MAFFT multiple sequence alignment software version 7: improvements in performance and usability, Mol Biol Evol, 30, 4, 772-80">Katoh and Standley 2013</a>) and trimmed by trimAl v1.4.rev15 with the default parameter settings (<a href="https://www.ncbi.nlm.nih.gov/pubmed/19505945" target="ictvref" title=" Capella-Gutiérrez et al., 2009, trimAl: a tool for automated alignment trimming in large-scale phylogenetic analyses, Bioinformatics, 25, 15, 1972-3">Capella-Gutiérrez <i>et al.,</i> 2009</a>). The phylogeny was inferred by IQ-TREE v.2.0.3 with the best fit model LG+G4 (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32011700" target="ictvref" title=" Minh et al., 2020, IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era, Mol Biol Evol, 37, 5, 1530-4">Minh <i>et al.,</i> 2020</a>). The confidence was analyzed by the ultrafast bootstrap method in 1000 replicates. The tree was constructed using MEGA X (<a href="https://www.ncbi.nlm.nih.gov/pubmed/29722887" target="ictvref" title=" Kumar et al., 2018, MEGA X: Molecular Evolutionary Genetics Analysis across Computing Platforms, Mol Biol Evol, 35, 6, 1547-9">Kumar <i>et al.,</i> 2018</a>).</p> </td> </tr> </tbody> </table> <h2>Relationships with other taxa</h2> <p>Hadakavirids are most closely related to members of the family <em>Polymycoviridae </em>(<a href="https://www.ncbi.nlm.nih.gov/pubmed/35639592" target="ictvref" title=" Kotta-Loizou et al., 2022, ICTV Virus Taxonomy Profile: Polymycoviridae 2022, J Gen Virol, 103, 5, ">Kotta-Loizou <i>et al.,</i> 2022</a>), based on phylogenetic analyses of the three conserved proteins (Hadaka/Polymyco-P1, P2 and P3 in Figure 2.<em>Hadakaviridae</em>) (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/34313859" target="ictvref" title=" Khan et al., 2021, A second capsidless hadakavirus strain with 10 positive-sense single-stranded RNA genomic segments from Fusarium nygamai, Arch Virol, 166, 10, 2711-22">Khan <i>et al.,</i> 2021</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/35435725" target="ictvref" title=" Fu et al., 2022, A novel heptasegmented positive-sense single-stranded RNA virus from the phytopathogenic fungus Colletotrichum fructicola, J Virol, 96, 9, e0031822">Fu <i>et al.,</i> 2022</a>). The largest difference between hadakavirids and polymycovirids is the absence or presence of the genome segment encoding a proline-alanine-serine-rich protein (PASrp). PASrp is known to physically associate with the dsRNA genome of polymycovirids (<a href="https://www.ncbi.nlm.nih.gov/pubmed/26139522" target="ictvref" title=" Kanhayuwa et al., 2015, A novel mycovirus from Aspergillus fumigatus contains four unique dsRNAs as its genome and is infectious as dsRNA, Proc Natl Acad Sci USA, 112, 29, 9100-5">Kanhayuwa <i>et al.,</i> 2015</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/33193262" target="ictvref" title=" Sato et al., 2020, Molecular characterization of a novel polymycovirus from Penicillium janthinellum with a focus on its genome-associated PASrp, Front Microbiol, 11, 592789">Sato <i>et al.,</i> 2020a</a>) and so gives polymycovirids an intermediate position between encapsidated and capsidless RNA viruses. Unlike polymycovirids and conventionally encapsidated viruses, HadV1 cannot be pelleted by ultracentrifugation and is susceptible to exogenously added ribonuclease A in host tissue homogenates (<a href="https://www.ncbi.nlm.nih.gov/pubmed/33193262" target="ictvref" title=" Sato et al., 2020, Molecular characterization of a novel polymycovirus from Penicillium janthinellum with a focus on its genome-associated PASrp, Front Microbiol, 11, 592789">Sato <i>et al.,</i> 2020a</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>).</p> <p>Based on the conserved RdRP motifs, hadakavirids show phylogenetic affinity to (+) RNA viruses, such as the members of the family <em>Astroviridae</em> and <em>Caliciviridae</em>, and is therefore classified in the phylum <em>Pisuviricota</em> (extended “picornavirus supergroup”) (<a href="https://www.ncbi.nlm.nih.gov/pubmed/32457242" target="ictvref" title=" Sato et al., 2020, Hadaka virus 1: a capsidless eleven-segmented positive-sense single-stranded RNA virus from a phytopathogenic fungus, Fusarium oxysporum, MBio, 11, 3, e00450-20">Sato <i>et al.,</i> 2020b</a>, <a href="https://www.ncbi.nlm.nih.gov/pubmed/35435725" target="ictvref" title=" Fu et al., 2022, A novel heptasegmented positive-sense single-stranded RNA virus from the phytopathogenic fungus Colletotrichum fructicola, J Virol, 96, 9, e0031822">Fu <i>et al.,</i> 2022</a>) along with other capsidless (+) RNA virus taxa such as the family <em>Hypoviridae</em> (<a href="https://www.ncbi.nlm.nih.gov/pubmed/29589826" target="ictvref" title=" Suzuki et al., 2018, ICTV Virus Taxonomy Profile: Hypoviridae, J Gen Virol, 99, 5, 615-6">Suzuki <i>et al.,</i> 2018</a>). The catalytic core amino acid residues of the RdRPs of hadakavirids and polymycovirids are “GDNQ”, which is a hallmark of some negative-sense RNA viruses in the order <em>Mononegavirales</em>, whereas the enzymes encoded by most members of the phylum <em>Pisuviricota</em> are characterized by a “GDD” triplet.</p> </div> </div> </div> </div> </div> </div></div> </div> </div> </div> </div> </div> </div> </article> </div> </div> </section> </div> </div> </div> </div> <div class="footers-container"> <div id="subfooter" class="clearfix subfooter region--shade-background region--no-separator region--no-block-paddings region--no-paddings"> <div class="container-fluid"> <div class="clearfix subfooter__container"> <div class="row align-items-center"> <div class="col-12 text-center"> <div class="clearfix subfooter__section subfooter-first"> <div class="region region-sub-footer-first"> <div id="block-ictvtitlesmall" class="contextual-region settings-tray-editable clearfix block 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