Nucleocytoviricota

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Nucleocytoviricota
Electron microscopic image of a mimivirus - journal.ppat.1000087.g007 crop.png

EM image of a virion (virus particle) of the
mimivirus , top left the 'Stargate' (opening
through which the DNA migrates into the host cell)

Systematics
Classification : Viruses
Area : Varidnaviria
Empire : Bamfordvirae
Phylum : Nucleocytoviricota
Taxonomic characteristics
Genome : dsDNA
Baltimore : Group 1
Scientific name
Nucleocytoviricota
Short name
NCLDV

The phylum of the nucleocytoviricota (formerly also English nucleocytoplasmic large DNA viruses , NCLDV ) comprises a heterogeneous group of mostly large dsDNA viruses that have a number of specific genes ( NCLDV core genes ) that common viruses lack. Founding members of the group (2001) are the families Asfarviridae , Iridoviridae and Poxviridae (smallpox viruses), as well as the Phycodnaviridae . Other members are the families Mimiviridae (synonymous with " Megaviridae "), Marseilleviridae , Ascoviridae . Proposed members are the pithoviruses , orpheoviruses and pandoraviruses , each with their own families if necessary, as well as many more. Possible candidates are dinodnavirus , " Medusa virus ", " Urceolovirus " and " Yaravirus ".

These members form - as has since been confirmed several times - a monophyletic kinship group, i. H. they share a common viral ancestor. It was therefore initially proposed on various occasions to raise this group as “ megavirales ” to the rank of a new virus order . After the International Committee on Taxonomy of Viruses (ICTV) had also allowed ranks above the order since 2018, such higher ranks were favored due to the high diversity of this group. This made it possible to set up the main groups of the NCLDV as orders or even classes. For example, Guglielmini et al. (2019) the extended family Mimiviridae as an order and suggested the now free term " Megavirales " for it. This development reached 2019/2020 with the Master Species List number 35 of the ICTV its preliminary conclusion with the official recognition of NCLDV as phylum Nucleocytoviricota (originally " Nucleocytoplasmaviricota ") with the classes Megaviricetes and Pokkesviricetes by ICTV in March 2020. The order of the extended Mimiviridae is now called Imitervirales .

Genome

The NCLDV genome is diverse and ranges in size from 100 to over 1500  kb (" Tupan virus "). Many NCLDV are giant viruses ( English: giant viruses, giruses ), which usually requires the 300 kb limit to be exceeded (for Yutin and Koonin of 500 kb). Conversely, all giant viruses (so far) are dsDNA viruses in the phylum NCLDV. The largest representatives form larger virus particles and have a larger genome than many small bacteria - the mimiviruses were initially even mistaken for ( parasitic ) bacteria, especially since they also show Gram staining . The late discovery of most giant viruses (numerous not until around 2003) was due to the fact that they got stuck in the filters (with a typical pore size of 0.2 μm) during the search for viruses, which were supposed to separate bacteria and protists from viruses easily visible clumps aggregate and also multiply more slowly.

Hosts

Although the NCLDV encode more proteins of their own than ordinary viruses (often hundreds instead of barely a dozen), they are host-specific. The hosts are exclusively complex-cellular organisms ( eukaryotes ): many protists (e.g. amoebas and algae ), but also vertebrates and insects. Originally six, now nine common ( homologous ) genes can be found in all NCLDV ( NCLDV core genes ), 177 further genes (status 2009) occur in at least two of the families. These include four genes encoding the DNA - replication concern and repair mechanisms: the DNA polymerase family B, the topoisomerase II A, the "flap" - endonuclease and the ring clamp protein ( proliferating cell nuclear antigen ), and the RNA -Polymerase II and the transcription factor II B. The genes of some NCLDV also contain introns .

Propagation cycle

Some of the NCLDV reproduce in whole or in part in the cell plasma (cytoplasm) of the eukaryotic host cells, others possibly go through a nuclear phase in the cell nucleus. Many NCLDV form after infection in the cytoplasm of their host, a microscopically visible production facility ( virus factory , English virus factory ) from. In some of these there are other viruses that use their synthesis apparatus for their own reproduction and are therefore called virophages ('virus eaters'). The first discovered case of this kind was Sputnik virus 1 with the NCLDV Mama virus , more Virophagen like Zamilon 'also attacked representatives of mimiviridae . Viruses are traditionally and mostly still not viewed as living beings, but the services provided by NCLDV reduce the gap to the living world. The fact that virions do not have a metabolism based on ATP as an “energy currency” remains unchanged .

Horizontal gene transfer

Massive horizontal gene transfer in both directions (AtoV, amoeba on virus and vice versa VtoA, virus on amoeba) can be detected between the giant viruses as viral endocytobionts (organisms that live or multiply in the cells of other organisms) and their often amoeboid hosts.

Didier Raoult's team interpreted the fact that some of the genes distributed among the NCLDV were not found to correspond to them in cellular organisms as an indication of a fourth domain of life, whose representatives have become extinct except for the parasitic NCLDV. This Forth Branch of Life hypothesis is no longer tenable since several genes distributed in NCLDV groups have been shown to originate from widely spaced locations in the tree of life, mainly from different eukaryotes, but also from bacteria. However, the discovery of each new family of giant viruses spurred the discussion anew, as the example of the Medusaviridae shows.

A summary of this discussion can be found in Traci Watson (2019).

In this context it was even proven that there is not only a gene transfer between the amoeboid hosts and the giant viruses as viral endocytobionts (VtoA / AtoV), but even between the viruses and bacterial endocytobionts that are present at the same time. Maybe this could be the odd match in the defense system CRISPR the bacteria to viruses (ie bacteriophages ) on one hand and the defense system MIMIVIRE the Mimiviren against parasitic satellite viruses (ie Virophage ) on the other hand explain.

Systematics

Internal systematics and research history

Despite their diversity, the NCLDV form a family group ( taxon ), as confirmed several times , whereby the giant viruses seem to be descended from (different) groups of smaller NCLDV, instead of the other way around.

  • " Irido-Ascoviridae ": Several studies since 2000 support the assumption that the Ascoviridae evolved from the Iridoviridae , but it could also be the other way around. It is also assumed that the genus Ichnovirus (family Polydnaviridae ) developed from the Ascoviridae .
  • Dinodnavirus : Investigations of the genome of Heterocapsa circularisquama DNA virus 01 (genus Dinodnavirus ) showed in 2009 that this genus, originally part of the Phycodnaviridae family, belongs to the Asfarviridae family.
  • " Urceolovirus ": Another species of giant DNA virus, " Urceolovirus corneum " ("KLaHel"), first reported in 2015, could be a member of the NCLDV.
  • New toilet viruses: In April 2017, reports were made about the discovery of four new giant viruses in the Klosterneuburg sewage treatment plant . Michael Wagner (Department of Microbiology and Ecosystem Research at the University of Vienna), Holger Daims, Matthias Horn and Frederik Schulz (all University of Vienna) published about these toilet new viruses (proposed subfamily of the Mimiviridae ) in Science .
  • " Orpheovirus ": In January 2018 Julien Andreani and colleagues reported about a new giant virus orpheovirus . According to the authors' suggestion, “ Orpheovirus ” should be placed in its own family “ Orpheoviridae ”, closely related to the proposed family Pithoviridae of the pithoviruses.
  • " Meelsvirus ": In September 2018, Shinn and Bullard found a giant virus (" Meelsvirus "), which infects the arrowworm Adhesisagitta hispida , replicates in the nucleus and envelops its virions with a length of 1.25  μm, when they again analyzed electron microscope images from the 1980s are. A better classification is not yet possible due to a lack of genome data.
Molecular phylogenetic analysis of NCLDV members using the maximum likelihood method . As of March 2017, source: MEGA7,
• Mimiviridae is to be understood in the broader sense (order Imitervirales ),
• The Asfar and Poxviridae form a clade (class Pokkesviricetes ),
• The Ascovirdae are paraphyletic and form a clade with the Iridoviridae (“ Irido -Ascoviridae ")
Size comparison of some giant viruses with HIV and a bacterium ( E. Coli )
  • " Medusavirus ": A proposal for another family " Medusaviridae " can be found at the CNRS (2018). This group could branch off basal from the NCLDV and thus perhaps form an additional fourth branch, or be located in the vicinity of the " Mollivirus ".
  • " Choanovirus ": In September 2019, David M. Needhal, Alexandra Z. Worden et al. about two species (1 and 2) of another new genus " Choanovirus " of the extended mimiviruses (now imitervirales ). The closest relative could be the " Aureococcus anophagefferens virus " ("AaV").
  • " Klothoviridae ": Also in 2019, Roxane-Marie Barthélémy et al. a family " Klothoviridae " with type species " Klothovirus casanovai " and another species " Megaklothovirus horridgei ", which would represent a new size record with 2.5-3.1 μm and 4 μm. Similar to the Arenaviridae , ribosomes were found in the virus particles. Since no genome data are currently available, a more precise classification of the family is not yet possible.
  • " Mininucleoviridae ": In January 2020, Subramaniam et al. Another family " Mininucleoviridae " of giant viruses proposed by the NCLDV, whose members parasitize crustaceans (Crustacea). According to the proposal, the members of this family include “ Carcinus maenas virus 1 ” (CmV1, alias “ Herpes-like virus ”, HLV), “ Dikerogammarus haemobaphes virus 1 ” (DhV1, alias “ Dikerogammarus haemobaphes bi-facieslike virus ”, DhbflV) and " Panulirus argus virus 1 " (PaV1). The family apparently belongs to the Pitho-Irido-Marseille branch of the NCLDV (now order Pimascovirales ).
  • “ Yaravirus ”: Also in January 2020, a preprint by Paulo VM Boratto, Bernard La Scola, Jônatas S. Abrahão et al. , in which they reported on a DNA virus with genes that sometimes differ greatly from the other NCLDV, for which they propose the name " Yaravirus brasiliensis ". The affiliation to the NCLDV and also to the Varidnaviria area is initially unclear.

System according to ICTV 2020

In March 2020, the ICTV followed the proposal by Koonin et al. (2015 and 2019) followed an internal system of the NCLDV, in which an extended family of the Asfarviridae (now order Asfuvirales ) form a sister group of the Poxviridae (with order Chitovirales ). From Koonin et al. (2015 and 2019) comes the so acceptable division into three main groups (or branches English branches ) as follows:

  • Phylum Nucleocytoviricota ( Nucleocytoplasmic large DNA viruses (NCLDV) , original proposal " Nucleocytoplasmaviricota ", replaces older proposals as the order " Megavirales " see left )
  • Branch 1:
  • Branch 2: order Pimascovirales (contraction of P itho , I rido , M arseille and Asco , formerly also called MAPI super clade or PMI group )
  • Branch 3:

The candidates according to Schulz et al. (2018), Rolland et al. (2019) and Subramaniam (2020), unless they belong to the extended Mimiviridae (order Imitervirales ). The candidates Dinodnavirus , “ Medusaviridae ”, “ Urceolovirus ” and “ Yaravirus ” have not yet been taken into account by the ICTV, which is in line with the proposal by Koonin et al. from April 2019.

Guglielmini et al. (2019) suggested the division into two super shops as follows:

  • MAPI super clade (branch 2 from above: pimascovirales )
  • PAM super clade (corresponds roughly to branches 1 and 3 (only Asfuvirales ))

The Poxviridae (smallpox viruses) are not included in this proposal.

Following an earlier suggestion by Schulz et al. (2018), Fig. 2, the Poxviridae, on the other hand, are basal in the NCLDV, the (extended) Asfarviridae were located in the branch of the Marseilleviridae . In contrast to Koonin et al. (2015 and 2019) the giant viruses still appeared here as in Guglielmini et al. monophyletic . However, the ICTV 2020 is based on the results of Koonin, Yutin, Bäckström, Ettema et al. followed after giant viruses developed several times from simple precursors within the NCLDV:

According to Clara Rolland et al. (2019), Fig. 2, the imitervirales form a common clade with the Pokkesviricetes , sister groups are the algavirales and pimascovirales on par. The Megaviricetes - although officially confirmed as a taxon by the ICTV in March 2020 - do not seem to form a clade as a sister group of the Pokkesviricetes .

According to Disa Bäckström et al. (2019), Fig. 1, seem - in contrast to Fig. 6 - the Pokkesviricetes similar to Schulz et al. (2018) not to form a common clade, the Poxviridae (or Chitovirales ) are basal, only then do the Asfarviridae (or Asfuvirales ) branch off from the rest of the NCLDV.

Other family trees based on neighbor-joining and maximum likelihood can be found among others. a. in the following works:

  • William H. Wilson et al. (2017), Fig. 2
  • Claire Bertelli et al. (2017), Fig. 4
  • Disa Bäckström (2018), Fig. 2
  • Disa Bäckström et al. (2019), Figs. 1 and 6
  • Sailen Barik (2018), Fig. 1

External system

The organization of the genome and the DNA replication mechanism create a phylogenetic relationship close between Rudi viruses (order ligamenvirales : rudivirus ) and large eukaryalen DNA viruses (NCLDVs) such as the African swine fever virus ( African swine fever virus , Asfarviridae ) Chloroviren ( Chlorella virus , Phycodnaviridae ) and smallpox viruses ( Orthopoxvirus , Poxviridae ).

Koonin et al. (2015, 2019) suspect the origin of the NCLDV in the Tectiviridae , icosahedral tailless ssDNA bacteriophages , according to ICTV also in the kingdom of Bamfordvirae - in contrast to the herpes viruses (herpes viruses), which are believed to originate from the tailed caudovirals . According to this proposal, the development proceeded via or by means of Polintoviruses ( Polintons , also called Mavericks: large DNA transposons which encode viral proteins, but also frequently occur in eukaryotic genomes). The development of adenoviruses ( Adenoviridae ) and bidnaviruses ( Bidnaviridae ) as well as of virophages (of the Lavidaviridae family ) was, it is assumed, initiated by the Polintons. The taxonomic classifications made by the ICTV in March 2020 take this into account: they organize

literature

  • Natalya Yutin, Yuri I. Wolf, Eugene V. Koonin: Origin of giant viruses from smaller DNA viruses not from a fourth domain of cellular life. In: Virology. 2014, doi: 10.1016 / j.virol.2014.06.032 , PMC 4325995 (free full text).
  • N. Yutin, P. Colson, D. Raoult, EV Koonin: Mimiviridae: clusters of orthologous genes, reconstruction of gene repertoire evolution and proposed expansion of the giant virus family. In: Virol J. 10, April 4, 2013, p. 106, PMID 23557328

Individual evidence

  1. ↑ Jan Diesend, Janis Kruse, Monica Hagedorn, Christian Hammann: Amoebae, Giant Viruses, and Virophages Make Up a Complex, Multilayered Threesome , in: Frontiers in Cellular and Infection Microbiology January 7, 2018, [doi: 10.3389 / fcimb.2017.00527] , via ResearchGate , Fig. 1 (“ Megavirals ” in the sense of NCLDV and NCLDV itself are not quite correctly referred to as “family” in this work, what is meant is “group”).
  2. ↑ a b c d e f g ICTV: ICTV Master Species List 2019.v1 , New MSL including all taxa updates since the 2018b release, March 2020 (MSL # 35)
  3. ↑ a b c Koonin EV, Dolja VV, Krupovic M, Varsani A, Wolf YI, Yutin N, Zerbini M, Kuhn JH: Create a megataxonomic framework for DNA viruses encoding vertical jelly roll-type major capsid proteins filling all principal taxonomic ranks . ICTV Proposal 2019.003G, April – July 2019
  4. ↑ Natalya Yutin, Yuri I. Wolf, Eugene V. Koonin: Origin of giant viruses from smaller DNA viruses not from a fourth domain of cellular life. In: Virology. 2014: " The 'Megavirales' unite 7 families of viruses ... ".
  5. ↑ a b c d e Genki Yoshikawa, Romain Blanc-Mathieu, Chihong Song, Yoko Kayama, Tomohiro Mochizuki, Kazuyoshi Murata, Hiroyuki Ogata, Masaharu Takemura: Medusavirus, a novel large DNA virus discovered from hot spring water . In: Journal of Virology . 93, No. 8, 2019. doi : 10.1128 / JVI.02130-18 . PMID 30728258 . Retrieved July 2, 2019. PDF
  6. ↑ a b Daniel Lingenhöhl: Giant virus turns host to stone , on: Spektrum.de of March 8, 2019
  7. ↑ a b Center national de la recherche scientifique: List of the main “giant” viruses known as of today (April, 18th, 2018) , Université Aix Marseille, April 18, 2018. According to Masaharu Takemura et al. : Giant Viruses Isolated from Japanese Aquatic Environments, Tokyo University of Science, 3rd Ringberg Symposium on Giant Virus Biology , 19. – 22. November 2017 (unpublished)
  8. ↑ a b Center national de la recherche scientifique: List of the main “giant” viruses known as of today (March 2019) , Université Aix Marseille, March 2019.
  9. ↑ a b Yetunde Adegboye, Mazen Atia, Anna M. Sigmund, Anne Krueger, Carsten Kirschning et al. : Abundance and Immunogenicity of Two Giant Viruses Namely Pithovirus lacustris and Urceolovirus corneum ( PDF ), Conference Paper, 14th International Conference on Microbial Interactions & Microbial Ecology, Vienna (Austria), August 2019
  10. ↑ a b Paulo VM Boratto, Graziel P. Oliveira, Talita B. Machado, Ana Cláudia SP Andrade, Jean-Pierre Baudoin, Thomas Klose, Frederik Schulz, Saïd Azza, Philippe Decloquement, Eric Chabrière, Philippe Colson, Anthony Levasseur, Bernard La Scola, Jônatas S. Abrahão: A mysterious 80 nm amoeba virus with a near-complete “ORFan genome” challenges the classification of DNA viruses , on: bioRxiv, January 28, 2020, bioRxiv : 10.1101 / 2020.01.28.923185v1 ( Preprint - Full text ) doi: 10.1101 / 2020.01.28.923185
  11. ↑ Lakshminarayan M. Iyer, L. Aravind, Eugene V. Koonin: Common Origin of Four Diverse Families of Large Eukaryotic DNA Viruses. In: Journal of Virology . 75, 2001, pp. 11720-11734, doi: 10.1128 / JVI.75.23.11720-11734.2001 , PMC 114758 (free full text).
  12. ↑ a b c d e f g Eugene V. Koonin, Natalya Yutin: Evolution of the Large Nucleocytoplasmatic DNA Viruses of Eukaryotes and Convergent Origins of Viral Gigantism , in: Advances in Virus research, Volume 103, AP January 21, 2019, doi: 10.1016 / bs.aivir.2018.09.002 , pp. 167-202. The toilet viruses are partially missed as toilet viruses.
  13. ^ A b c Eugene V. Koonin, Valerian V. Dolja, Mart Krupovic: Origins and evolution of viruses of eukaryotes: The ultimate modularity , in: Virology from May 2015; 479-480. 2-25, Epub March 12, 2015, PMC 5898234 (free full text), PMID 25771806
  14. ↑ Natalya Yutin, Yuri I. Wolf, Eugene V. Koonin: Origin of giant viruses from smaller DNA viruses not from a fourth domain of cellular life. In: Virology. October 2014, doi: 10.1016 / j.virol.2014.06.032 , PMID 25042053 : " ... we refer to this major group of viruses as 'Megavirales' to signal our support of this amendment to virus taxonomy ... ", a suggestion from the working group around Didier Raoult, see P. Colson, X. de Lamballerie, G. Fournous, D. Raoult: Reclassification of giant viruses composing a fourth domain of life in the new order Megavirales. In: Intervirology. 55, 2012, pp. 321-332, doi: 10.1159 / 000336562 , PMID 22508375 .
  15. ↑ ICTV Executive Committee: The new scope of virus taxonomy: partitioning the virosphere into 15 hierarchical ranks , in: Nature Microbiology 5, pp. 668–674, April 27, 2020, doi: 10.1038 / s41564-020-0709-x , PDF . Status: January 2020
  16. ↑ Nadja Podbregar: A family tree for the virosphere , on: scinexx.de from April 29, 2020. As of January 2020
  17. ↑ a b c d Julien Guglielmini, Anthony C. Woo, Mart Krupovic, Patrick Forterre, Morgan Gaia: Diversification of giant and large eukaryotic dsDNnA viruses predated the origin of modern eukaryotes , in: PNAS 116 (39), 10./24. September 2019, pp. 19585-19592, doi: 10.1073 / pnas.1912006116 , PMID 31506349 , Fig. 2
  18. ↑ N. Yutin, YI Wolf, EV Koonin: Origin of giant viruses from smaller DNA viruses not from a fourth domain of cellular life. In: Virology. 2014: " All giant viruses contain a set of core genes that define ... the proposed order Megavirales ".
  19. ↑ a b James L. Van Etten, Leslie C. Lane, David D. Dunigan: DNA Viruses: The Really Big Ones (Giruses) . Annu. Rev. Microbiol. 64, 2010, pp. 83-99, doi: 10.1146 / annurev.micro.112408.134338 ( full text free for personal use ).
  20. ↑ Natalya Yutin, Philippe Colson, Didier Raoult, Eugene V. Koonin: Mimiviridae: clusters of orthologous genes, reconstruction of gene repertoire evolution and proposed expansion of the giant virus family. In: Virol J. 10, 2013, p. 106, doi: 10.1186 / 1743-422X-10-106 . PMID 23557328 (free full text).
  21. ^ A b Anthony Levasseur, Meriem Bekliz, Eric Chabrière, Pierre Pontarotti, Bernard La Scola and Didier Raoult: MIMIVIRE is a defense system in mimivirus that confers resistance to virophage. In: Nature. 2016, doi: 10.1038 / nature17146 (free full text).
  22. ↑ d. H. intracellular symbiont or parasite
  23. ↑ Natalya Yutin, Yuri I. Wolf, Eugene V. Koonin: Origin of giant viruses from smaller DNA viruses not from a fourth domain of cellular life. In: Virology. 2014, Figs. 1 to 4.
  24. ↑ a b c d e Disa Bäckström, Natalya Yutin, Steffen L. Jørgensen, Jennah Dharamshi, Felix Homa, Katarzyna Zaremba-Niedwiedzka, Anja Spang, Yuri I. Wolf, Eugene V. Koonin, Thijs J. G. Ettema; Richard P. Novick (Ed.): Virus Genomes from Deep Sea Sediments Expand the Ocean Megavirome and Support Independent Origins of Viral Gigantism , in: mBio Vol. 10, No. 2, March – April 2019, pp. E02497-18, PDF , doi: 10.1128 / mBio.02497-18 , PMC 6401483 (free full text), PMID 30837339 , ResearchGate . Note: In Fig. 1 it must be Megamimivirinae (or Megavirinae ) instead of Mimiviridae . This subfamily, together with the Klosneuvirinae and the cafeteria viruses, forms the family Mimiviridae s. s.
  25. ↑ Traci Watson: The trickster microbes that are shaking up the tree of life , in: Nature from May 14, 2019 (English), Trickser bacteria shake the family tree of life , in: Spektrum.de from June 20, 2019 (German) - the term 'bacteria' is not entirely correct; the microbes considered are archaea or (according to some researchers) at least proto- eukaryotes different from bacteria .
  26. ↑ Patrick L. Scheid: Free-Living Amoebae and Their Multiple Impacts on Environmental Health , in: Reference Module in Earth Systems and Environmental Sciences, February 27, 2018, doi: 10.1016 / B978-0-12-409548-9.10969-8 , here: text following Fig. 8 (right column)
  27. ↑ Takashi Yamada: Giant viruses in the environment: Their origins and evolution , in: Curr Opin Virol .; July 2011; 1 (1); Pp. 58-62, doi: 10.1016 / j.coviro.2011.05.008 , PMID 22440568
  28. ↑ N. Yutin, YI Wolf, EV Koonin: Origin of giant viruses from smaller DNA viruses not from a fourth domain of cellular life. In: Virology. 2014, Figs. 5 and 6.
  29. ↑ K. Stasiak, MV Demattei, BA Federici, Y. Bigot: Phylogenetic position of the Diadromus pulchellus ascovirus DNA polymerase among viruses with large double-stranded DNA genomes . In: The Journal of General Virology . tape 81 , Pt 12, December 2000, p. 3059-72 , doi : 10.1099 / 0022-1317-81-12-3059 , PMID 11086137 .
  30. ↑ K. Stasiak, S. Renault, MV Demattei, Y. Bigot, BA Federici: Evidence for the evolution of ascoviruses from iridoviruses . In: The Journal of General Virology . tape 84 , Pt 11, November 2003, p. 2999-3009 , doi : 10.1099 / vir.0.19290-0 , PMID 14573805 .
  31. ↑ a b B. A. Federici, DK Bideshi, Y. Tan, T. Spears, Y. Bigot: Ascoviruses: superb manipulators of apoptosis for viral replication and transmission . In: Current Topics in Microbiology and Immunology . tape 328 , 2009, ISBN 978-3-540-68617-0 , pp. 171-96 , doi : 10.1007 / 978-3-540-68618-7_5 , PMID 19216438 .
  32. ↑ B. Piégu, S. Asgari, D. Bideshi, BA Federici, Y. Bigot: Evolutionary relationships of iridoviruses and divergence of ascoviruses from invertebrate iridoviruses in the superfamily Megavirales . In: Molecular Phylogenetics and Evolution . tape 84 , March 2015, p. 44-52 , doi : 10.1016 / j.ympev.2014.12.013 , PMID 25562178 .
  33. ↑ dsDNA Viruses> Ascoviridae , on: ICTV online, December 2016 (here: Fig. 2)
  34. ↑ Fenner's Veterinary Virology (Fifth Edition): Chapter 8 - Asfarviridae and Iridoviridae , online November 4, 2016, pp. 175–188, doi: 10.1016 / B978-0-12-800946-8.00008-8 (here: Fig. 1 )
  35. ↑ a b c Hiroyuki Ogata, Kensuke Toyoda, Yuji Tomaru, Natsuko Nakayama, Yoko Shirai, Jean-Michel Claverie and Keizo Nagasaki: Remarkable sequence similarity between the dinoflagellate-infecting marine girus and the terrestrial pathogen African swine fever virus. In: Virol J. 6, 2009, doi: 10.1186 / 1743-422X-6-178 , p. 178.
  36. ↑ NCBI: Urceolovirus corneum (species)
  37. ↑ SAMN06480674 - U. corneum nucleotide , on: BioSamples, European Bioinformatics Institute (GB)
  38. ↑ Urceolovirus corneum (species) , on: DDBJ DNA Data bank of Japan
  39. ↑ Researchers puzzling over giant viruses from sewage treatment plant orf.at, April 6, 2017, accessed April 6, 2017.
  40. ↑ a b c d Julien Andreani, Jacques Y.B. Khalil, Emeline Baptiste, Issam Hasni, Caroline Michelle, Didier Raoult, Anthony Levasseur, Bernard La Scola: Orpheovirus IHUMI-LCC2: A New Virus among the Giant Viruses . In: Frontiers in Microbiology . tape 8 , January 22, 2018, ISSN  1664-302X , doi : 10.3389 / fmicb.2017.02643 ( frontiersin.org [PDF]).
  41. ↑ G. L. Shinn, B. L. Bullard; Carmen San Martin (Ed.): Ultrastructure of Meelsvirus: A nuclear virus of arrow worms (phylum Chaetognatha) producing giant "tailed" virions . In: PLOS ONE . 13, No. 9, September 19, 2018, p. E0203282. bibcode : 2018PLoSO..1303282S . doi : 10.1371 / journal.pone.0203282 . PMID 30231047 . PMC 6145532 (free full text).
  42. ^ A b Roxane-Marie Barthélémy, Eric Faure, Taichiro Goto: Serendipitous Discovery in a Marine Invertebrate (Phylum Chaetognatha) of the Longest Giant Viruses Reported till Date , in: Biology 2019, Abstract
  43. ↑ NCBI: Sylvanvirus (species)
  44. ↑ NCBI: Solivirus (species)
  45. ↑ NCBI: Solum virus (species)
  46. ↑ a b c Frederik Schulz, Lauren Alteio, Danielle Goudeau, Elizabeth M. Ryan, Feiqiao B. Yu, Rex R. Malmstrom, Jeffrey Blanchard, Tanja Woyke: Hidden diversity of soil giant viruses , in: Nature Communicationsvolume 9, Article number: 4881 (2018) of November 19, 2018, doi: 10.1038 / s41467-018-07335-2
  47. ↑ Jan Osterkamp: Virology: Giant viruses are more widespread than expected , on: Spektrum.de from November 20, 2018.
  48. ↑ Sudhir Kumar, Glen Stecher, Koichiro Tamura: MEGA7: Molecular Evolutionary Genetics Analysis Version 7.0 for Bigger Datasets . In: Molecular Biology and Evolution . 33, No. 7, 2016, pp. 1870–1874. doi : 10.1093 / molbev / msw054 . PMID 27004904 .
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