The genome of Diabrotica undecimpunctata virus 2 (DuV2), a positive-sense, single-stranded RNA virus identified from the southern corn root- worm transcriptome, comprises 5,313 nucleotides, including a short poly(A) tail. The two open reading frames encode a nonstructural polyprotein (p156) and a putative capsid protein (p25)

Bonning, Bryony C.

Darlington, Molly

Liu, Sijun

Valencia-Jiménez, Arnubio

Vélez, Ana M.

English

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University of Nebraska - Lincoln 
DigitalCommons@University of Nebraska - Lincoln 
Faculty Publications: Department of 
Entomology Entomology, Department of 
2020 
Diabrotica undecimpunctata virus 2, a Novel Small RNA Virus 
Discovered from Southern Corn Rootworm, Diabrotica 
undecimpunctata howardi Barber (Coleoptera: Chrysomelidae) 
Sijun Liu 
Arnubio Valencia-Jiménez 
Molly Darlington 
Ana M. Vélez 
Bryony C. Bonning 
Follow this and additional works at: https://digitalcommons.unl.edu/entomologyfacpub 
 Part of the Entomology Commons 
This Article is brought to you for free and open access by the Entomology, Department of at 
DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Faculty Publications: 
Department of Entomology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. 
Diabrotica undecimpunctata virus 2, a Novel Small RNA Virus
Discovered from Southern Corn Rootworm, Diabrotica
undecimpunctata howardi Barber (Coleoptera: Chrysomelidae)
Sijun Liu,a Arnubio Valencia-Jiménez,b,c Molly Darlington,c Ana M. Vélez,c Bryony C. Bonningd
aDepartment of Entomology, Iowa State University, Ames, Iowa, USA
bDepartamento de Producción Agropecuaria, Universidad de Caldas, Manizales, Colombia
cDepartment of Entomology, University of Nebraska—Lincoln, Lincoln, Nebraska, USA
dDepartment of Entomology and Nematology, University of Florida, Gainesville, Florida, USA
ABSTRACT The genome of Diabrotica undecimpunctata virus 2 (DuV2), a putative
positive-sense, single-stranded RNA virus identified from the southern corn root-
worm transcriptome, comprises 5,313 nucleotides, including a short poly(A) tail. The
two open reading frames encode a nonstructural polyprotein (p156) and a putative
capsid protein (p25).
The southern corn rootworm (SCR), Diabrotica undecimpunctata howardi Barber, isone of three corn rootworm species that cause significant damage to agricultural
and horticultural crops (1). Several small RNA viruses have been identified from the
western corn rootworm, Diabrotica virgifera virgifera (2–4), but only one has been
described from SCR to date (5). Here, we report on the genome of a second RNA virus
identified from the transcriptome of SCR.
Eggs, larvae (first, second, and third instar), pupae, and adult (male and female) SCR
were supplied by Crop Characteristics (Farmington, MN). The RNeasy minikit (Qiagen,
Valencia, CA, USA) was used for isolation of total RNA from three biological replicates
of 100 eggs, 50 first-instar larvae, 5 each of second- and third-instar larvae, 1 pupa, 1
adult female, and 1 adult male. cDNA libraries were constructed using the NEBNext
Ultra RNA library prep kit for Illumina (NEB) for each stage-specific sample with poly(A)
selection and paired-end sequencing with 150-nucleotide (nt) reads conducted on a
HiSeq 2500 platform (Illumina, San Diego, CA). One or two replicate sequence data sets
from each developmental stage were used for virus sequence discovery (5).
The Trinity assembler v2.6.6 (6) was used for assembly of high-quality reads. Default
parameters were used for all software unless otherwise noted. A putative viral contig,
with a short poly(A) tail, has 5,312 nt with a nucleotide composition of 37% GC
content. A total of 29,036 and 29,694 reads from two first-instar replicate data sets
(comprising 50.7 million and 48.1 million reads and quality scores of 37.5 and 37.6,
respectively) mapped to the viral contig with coverages of 819.8- and 838.3-fold,
respectively. The virus is provisionally named Diabrotica undecimpunctata virus 2
(DuV2). Sequences derived from DuV2 were observed in transcriptome data sets from
all developmental stages of SCR. The near-full-length sequences were assembled from
first-instar larvae, suggesting increased abundance and active replication of DuV2 in
larvae relative to other developmental stages. The 5= coding sequence located between
nt 156 and 4286 encodes a putative nonstructural protein of 1,376 amino acids (aa)
(156.1 kDa, p156), while the more 3= sequence between nt 4366 and 5025 encodes a
putative structural protein with 219 aa (25 kDa, p25). A conserved protein domain
search revealed that p156 contains three conserved domains, two overlapping viral
helicase domains (NCBI accession no. pfam01443, aa 598 to 826; cd17914, aa 593 to
Citation Liu S, Valencia-Jiménez A, Darlington
M, Vélez AM, Bonning BC. 2020. Diabrotica
undecimpunctata virus 2, a novel small RNA
virus discovered from southern corn rootworm,
Diabrotica undecimpunctata howardi Barber
(Coleoptera: Chrysomelidae). Microbiol Resour
Announc 9:e00380-20. https://doi.org/10.1128/
MRA.00380-20.
Editor Kenneth M. Stedman, Portland State
University
Copyright © 2020 Liu et al. This is an open-
access article distributed under the terms of
the Creative Commons Attribution 4.0
International license.
Address correspondence to Bryony C. Bonning,
bbonning@ufl.edu.
Received 10 April 2020
Accepted 28 May 2020
Published 25 June 2020
GENOME SEQUENCES
crossm
Volume 9 Issue 26 e00380-20 mra.asm.org 1
FIG 1 Phylogenetic analysis of DuV2 putative nonstructural protein p156 and putative structural protein p25. The replication-associated p156 is
homologous to proteins from viruses with various hosts, including plants, fungi, insects, and hepatitis E-like viruses. Sequences with similarity to the
capsid protein p25 were identified from viruses isolated from arthropods, although p25 also has a plant viral coat protein TMV domain. Viral proteins
used for construction of the phylogenetic tree were selected from BLASTp analyses. Sequences were aligned with MAFFT v7.4.07 (using the AUTO
strategy), and the maximum likelihood (ML) method was used for construction of the phylogenetic tree. ModelFinder implemented in IQ-TREE was
used to identify the best partitioning scheme and models. ML analysis was performed using IQ-TREE with 10,000 ultrafast bootstrap replicates (10).
Interactive Tree of Life (http://itol.embl.de) was used for visualization and annotation. The viruses and associated NCBI genome accession numbers
are Adelphocoris suturalis virus, YP_009336480.1; Agaricus bisporus virus 8, AQM49930.1; Agaricus bisporus virus 13, AQM49941.1; Bactrocera oleae,
XP_014085601.1; Bat bastrovirus, AWV67082.1; Beet necrotic yellow vein virus, ACA63016.1; Beet soil-borne mosaic virus, YP_009513207.1; Beihai
charybdis crab virus 1, YP_009333243.1; Burdock mottle virus, YP_008219063.1; Chara australis virus, AEJ33768.1 and AEJ33771.1; Culex pipiens
associated Tunisia virus, YP_009553258.1; Goji berry chlorosis virus, AYO99569.1; Hepatitis E virus, BAO47889.1; Hubei beny-like virus 1, APG77690.1;
Hubei virga-like virus 2, YP_009337413.1; Hubei virga-like virus 11, AVK59472.1 and YP_009337245.1; Lentinula edodes ssRNA mycovirus, BAS04359.1;
Mangifera indica latent virus, AMQ23297.1; Megastigmus ssRNA virus, QDZ71187.1; Red clover RNA virus 1, AVD98637.1; Riboviria sp. QDH86483.1; Rice
stripe necrosis virus, QAR21011.1; Rodent hepevirus, AVW79978.1; Sclerotium rolfsii beny-like virus 1, AZF86092.1; Wenling thamnaconus septentrionalis
hepevirus, AVM87557.1; Wheat stripe mosaic virus, AYD38100.1; and Zeugodacus cucurbitae; XP_011177481.1.
Liu et al.
Volume 9 Issue 26 e00380-20 mra.asm.org 2
726) (7), and one RNA-dependent RNA polymerase domain (RdRP_2; pfam00978). A
Tomato mosaic virus coat protein domain (TMV_coat domain pfam00721) was identified
in protein p25, supporting annotation as a capsid protein.
The highest BLASTp score for p156 (8) was Hubei beny-like virus 1 (GenBank
accession no. KX883789.1), an RNA virus isolated from dipteran insects (9) with 76%
sequence coverage and 49% amino acid identity. Proteins similar to p156 included
those from viruses of various host species (including fungi and plants) and soil samples
(Riboviria sp.) (Fig. 1). BLASTp analysis for p25 identified the putative capsid protein of
Beihai charybdis crab virus 1 (GenBank accession no. YP_009333243.1), which has a
genomic structure similar to that of DuV2.
Data availability. The genome sequence of DuV2 is available in GenBank under
accession no. MN646771. The raw reads for SRA accession no. SRX8018653 to
SRX8018656 are available at BioProject no. PRJNA615920.
ACKNOWLEDGMENTS
This research was supported by the USDA National Institute of Food and Agriculture,
Hatch project IOW05450; State of Iowa funds; University of Nebraska—Lincoln funds;
the 2016 DuPont Young Professor Grant awarded to A. M. Vélez; and the University of
Florida Institute of Food and Agricultural Sciences.
REFERENCES
1. Krysan JL. 1986. Introduction: biology, distribution, and identification of
pest Diabrotica, p 1–23. In Krysan JLM, Thomas A (ed), Methods for the
study of pest Diabrotica. Springer, New York, NY.
2. Liu S, Chen Y, Sappington TW, Bonning BC. 2017. Genome sequence of
a novel positive-sense, single-stranded RNA virus isolated from western
corn rootworm, Diabrotica virgifera virgifera LeConte. Genome Announc
5:e00366-17. https://doi.org/10.1128/genomeA.00366-17.
3. Liu S, Chen Y, Sappington TW, Bonning BC. 2017. Genome sequence of
Diabrotica virgifera virgifera virus 2, a novel small RNA virus of the
western corn rootworm, Diabrotica virgifera virgifera LeConte. Genome
Announc 5:e00365-17. https://doi.org/10.1128/genomeA.00365-17.
4. Liu S, Chen Y, Sappington TW, Bonning BC. 2017. Genome sequence of
the first coleopteran iflavirus isolated from western corn rootworm,
Diabrotica virgifera virgifera LeConte. Genome Announc 5:e01530-16.
https://doi.org/10.1128/genomeA.01530-16.
5. Liu S, Valencia-Jiménez A, Darlington M, Vélez AM, Bonning BC. 2020.
Genome sequence of a small RNA virus of the southern corn rootworm,
Diabrotica undecimpunctata howardi Barber (Coleoptera: Chrysomeli-
dae). Microbiol Resour Announc 9:e00379-20. https://doi.org/10.1128/
MRA.00379-20.
6. Haas BJ, Papanicolaou A, Yassour M, Grabherr M, Blood PD, Bowden
J, Couger MB, Eccles D, Li B, Lieber M, MacManes MD, Ott M, Orvis J,
Pochet N, Strozzi F, Weeks N, Westerman R, William T, Dewey CN,
Henschel R, Leduc RD, Friedman N, Regev A. 2013. De novo transcript
sequence reconstruction from RNA-seq using the Trinity platform for
reference generation and analysis. Nat Protoc 8:1494 –1512. https://
doi.org/10.1038/nprot.2013.084.
7. Marchler-Bauer A, Bo Y, Han L, He J, Lanczycki CJ, Lu S, Chitsaz F,
Derbyshire MK, Geer RC, Gonzales NR, Gwadz M, Hurwitz DI, Lu F,
Marchler GH, Song JS, Thanki N, Wang Z, Yamashita RA, Zhang D, Zheng
C, Geer LY, Bryant SH. 2017. CDD/SPARCLE: functional classification of
proteins via subfamily domain architectures. Nucleic Acids Res 45:
D200 –D203. https://doi.org/10.1093/nar/gkw1129.
8. Altschul SF, Wootton JC, Gertz EM, Agarwala R, Morgulis A, Schaffer AA,
Yu YK. 2005. Protein database searches using compositionally adjusted
substitution matrices. FEBS J 272:5101–5109. https://doi.org/10.1111/j
.1742-4658.2005.04945.x.
9. Shi M, Lin XD, Tian JH, Chen LJ, Chen X, Li CX, Qin XC, Li J, Cao JP, Eden
JS, Buchmann J, Wang W, Xu J, Holmes EC, Zhang YZ. 2016. Redefining
the invertebrate RNA virosphere. Nature 540:539 –543. https://doi.org/
10.1038/nature20167.
10. Hoang DT, Chernomor O, von Haeseler A, Minh BQ, Vinh LS. 2018.
UFBoot2: improving the ultrafast bootstrap approximation. Mol Biol Evol
35:518 –522. https://doi.org/10.1093/molbev/msx281.
Microbiology Resource Announcement
Volume 9 Issue 26 e00380-20 mra.asm.org 3


Diabrotica undecimpunctata virus 2, a Novel Small RNA Virus Discovered from Southern Corn Rootworm, Diabrotica undecimpunctata howardi Barber (Coleoptera: Chrysomelidae)

DigitalCommons@University of Nebraska

University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Faculty Publications: Department of Entomology Entomology, Department of 2020 Diabrotica undecimpunctata virus 2, a Novel Small RNA Virus Discovered from Southern Corn Rootworm, Diabrotica undecimpunctata howardi Barber (Coleoptera: Chrysomelidae) Sijun Liu Arnubio Valencia-Jiménez Molly Darlington Ana M. Vélez Bryony C. Bonning Follow this and additional works at: https://digitalcommons.unl.edu/entomologyfacpub  Part of the Entomology Commons This Article is brought to you for free and open access by the Entomology, Department of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Faculty Publications: Department of Entomology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Diabrotica undecimpunctata virus 2, a Novel Small RNA VirusDiscovered from Southern Corn Rootworm, Diabroticaundecimpunctata howardi Barber (Coleoptera: Chrysomelidae)Sijun Liu,a Arnubio Valencia-Jiménez,b,c Molly Darlington,c Ana M. Vélez,c Bryony C. BonningdaDepartment of Entomology, Iowa State University, Ames, Iowa, USAbDepartamento de Producción Agropecuaria, Universidad de Caldas, Manizales, ColombiacDepartment of Entomology, University of Nebraska—Lincoln, Lincoln, Nebraska, USAdDepartment of Entomology and Nematology, University of Florida, Gainesville, Florida, USAABSTRACT The genome of Diabrotica undecimpunctata virus 2 (DuV2), a putativepositive-sense, single-stranded RNA virus identified from the southern corn root-worm transcriptome, comprises 5,313 nucleotides, including a short poly(A) tail. Thetwo open reading frames encode a nonstructural polyprotein (p156) and a putativecapsid protein (p25).The southern corn rootworm (SCR), Diabrotica undecimpunctata howardi Barber, isone of three corn rootworm species that cause significant damage to agriculturaland horticultural crops (1). Several small RNA viruses have been identified from thewestern corn rootworm, Diabrotica virgifera virgifera (2–4), but only one has beendescribed from SCR to date (5). Here, we report on the genome of a second RNA virusidentified from the transcriptome of SCR.Eggs, larvae (first, second, and third instar), pupae, and adult (male and female) SCRwere supplied by Crop Characteristics (Farmington, MN). The RNeasy minikit (Qiagen,Valencia, CA, USA) was used for isolation of total RNA from three biological replicatesof 100 eggs, 50 first-instar larvae, 5 each of second- and third-instar larvae, 1 pupa, 1adult female, and 1 adult male. cDNA libraries were constructed using the NEBNextUltra RNA library prep kit for Illumina (NEB) for each stage-specific sample with poly(A)selection and paired-end sequencing with 150-nucleotide (nt) reads conducted on aHiSeq 2500 platform (Illumina, San Diego, CA). One or two replicate sequence data setsfrom each developmental stage were used for virus sequence discovery (5).The Trinity assembler v2.6.6 (6) was used for assembly of high-quality reads. Defaultparameters were used for all software unless otherwise noted. A putative viral contig,with a short poly(A) tail, has 5,312 nt with a nucleotide composition of 37% GCcontent. A total of 29,036 and 29,694 reads from two first-instar replicate data sets(comprising 50.7 million and 48.1 million reads and quality scores of 37.5 and 37.6,respectively) mapped to the viral contig with coverages of 819.8- and 838.3-fold,respectively. The virus is provisionally named Diabrotica undecimpunctata virus 2(DuV2). Sequences derived from DuV2 were observed in transcriptome data sets fromall developmental stages of SCR. The near-full-length sequences were assembled fromfirst-instar larvae, suggesting increased abundance and active replication of DuV2 inlarvae relative to other developmental stages. The 5= coding sequence located betweennt 156 and 4286 encodes a putative nonstructural protein of 1,376 amino acids (aa)(156.1 kDa, p156), while the more 3= sequence between nt 4366 and 5025 encodes aputative structural protein with 219 aa (25 kDa, p25). A conserved protein domainsearch revealed that p156 contains three conserved domains, two overlapping viralhelicase domains (NCBI accession no. pfam01443, aa 598 to 826; cd17914, aa 593 toCitation Liu S, Valencia-Jiménez A, DarlingtonM, Vélez AM, Bonning BC. 2020. Diabroticaundecimpunctata virus 2, a novel small RNAvirus discovered from southern corn rootworm,Diabrotica undecimpunctata howardi Barber(Coleoptera: Chrysomelidae). Microbiol ResourAnnounc 9:e00380-20. https://doi.org/10.1128/MRA.00380-20.Editor Kenneth M. Stedman, Portland StateUniversityCopyright © 2020 Liu et al. This is an open-access article distributed under the terms ofthe Creative Commons Attribution 4.0International license.Address correspondence to Bryony C. Bonning,bbonning@ufl.edu.Received 10 April 2020Accepted 28 May 2020Published 25 June 2020GENOME SEQUENCEScrossmVolume 9 Issue 26 e00380-20 mra.asm.org 1FIG 1 Phylogenetic analysis of DuV2 putative nonstructural protein p156 and putative structural protein p25. The replication-associated p156 ishomologous to proteins from viruses with various hosts, including plants, fungi, insects, and hepatitis E-like viruses. Sequences with similarity to thecapsid protein p25 were identified from viruses isolated from arthropods, although p25 also has a plant viral coat protein TMV domain. Viral proteinsused for construction of the phylogenetic tree were selected from BLASTp analyses. Sequences were aligned with MAFFT v7.4.07 (using the AUTOstrategy), and the maximum likelihood (ML) method was used for construction of the phylogenetic tree. ModelFinder implemented in IQ-TREE wasused to identify the best partitioning scheme and models. ML analysis was performed using IQ-TREE with 10,000 ultrafast bootstrap replicates (10).Interactive Tree of Life (http://itol.embl.de) was used for visualization and annotation. The viruses and associated NCBI genome accession numbersare Adelphocoris suturalis virus, YP_009336480.1; Agaricus bisporus virus 8, AQM49930.1; Agaricus bisporus virus 13, AQM49941.1; Bactrocera oleae,XP_014085601.1; Bat bastrovirus, AWV67082.1; Beet necrotic yellow vein virus, ACA63016.1; Beet soil-borne mosaic virus, YP_009513207.1; Beihaicharybdis crab virus 1, YP_009333243.1; Burdock mottle virus, YP_008219063.1; Chara australis virus, AEJ33768.1 and AEJ33771.1; Culex pipiensassociated Tunisia virus, YP_009553258.1; Goji berry chlorosis virus, AYO99569.1; Hepatitis E virus, BAO47889.1; Hubei beny-like virus 1, APG77690.1;Hubei virga-like virus 2, YP_009337413.1; Hubei virga-like virus 11, AVK59472.1 and YP_009337245.1; Lentinula edodes ssRNA mycovirus, BAS04359.1;Mangifera indica latent virus, AMQ23297.1;Megastigmus ssRNA virus, QDZ71187.1; Red clover RNA virus 1, AVD98637.1; Riboviria sp. QDH86483.1; Ricestripe necrosis virus, QAR21011.1; Rodent hepevirus, AVW79978.1; Sclerotium rolfsii beny-like virus 1, AZF86092.1;Wenling thamnaconus septentrionalishepevirus, AVM87557.1; Wheat stripe mosaic virus, AYD38100.1; and Zeugodacus cucurbitae; XP_011177481.1.Liu et al.Volume 9 Issue 26 e00380-20 mra.asm.org 2726) (7), and one RNA-dependent RNA polymerase domain (RdRP_2; pfam00978). ATomato mosaic virus coat protein domain (TMV_coat domain pfam00721) was identifiedin protein p25, supporting annotation as a capsid protein.The highest BLASTp score for p156 (8) was Hubei beny-like virus 1 (GenBankaccession no. KX883789.1), an RNA virus isolated from dipteran insects (9) with 76%sequence coverage and 49% amino acid identity. Proteins similar to p156 includedthose from viruses of various host species (including fungi and plants) and soil samples(Riboviria sp.) (Fig. 1). BLASTp analysis for p25 identified the putative capsid protein ofBeihai charybdis crab virus 1 (GenBank accession no. YP_009333243.1), which has agenomic structure similar to that of DuV2.Data availability. The genome sequence of DuV2 is available in GenBank underaccession no. MN646771. The raw reads for SRA accession no. SRX8018653 toSRX8018656 are available at BioProject no. PRJNA615920.ACKNOWLEDGMENTSThis research was supported by the USDA National Institute of Food and Agriculture,Hatch project IOW05450; State of Iowa funds; University of Nebraska—Lincoln funds;the 2016 DuPont Young Professor Grant awarded to A. M. Vélez; and the University ofFlorida Institute of Food and Agricultural Sciences.REFERENCES1. Krysan JL. 1986. Introduction: biology, distribution, and identification ofpest Diabrotica, p 1–23. In Krysan JLM, Thomas A (ed), Methods for thestudy of pest Diabrotica. Springer, New York, NY.2. Liu S, Chen Y, Sappington TW, Bonning BC. 2017. Genome sequence ofa novel positive-sense, single-stranded RNA virus isolated from westerncorn rootworm, Diabrotica virgifera virgifera LeConte. Genome Announc5:e00366-17. https://doi.org/10.1128/genomeA.00366-17.3. Liu S, Chen Y, Sappington TW, Bonning BC. 2017. Genome sequence ofDiabrotica virgifera virgifera virus 2, a novel small RNA virus of thewestern corn rootworm, Diabrotica virgifera virgifera LeConte. GenomeAnnounc 5:e00365-17. https://doi.org/10.1128/genomeA.00365-17.4. Liu S, Chen Y, Sappington TW, Bonning BC. 2017. Genome sequence ofthe first coleopteran iflavirus isolated from western corn rootworm,Diabrotica virgifera virgifera LeConte. Genome Announc 5:e01530-16.https://doi.org/10.1128/genomeA.01530-16.5. Liu S, Valencia-Jiménez A, Darlington M, Vélez AM, Bonning BC. 2020.Genome sequence of a small RNA virus of the southern corn rootworm,Diabrotica undecimpunctata howardi Barber (Coleoptera: Chrysomeli-dae). Microbiol Resour Announc 9:e00379-20. https://doi.org/10.1128/MRA.00379-20.6. Haas BJ, Papanicolaou A, Yassour M, Grabherr M, Blood PD, BowdenJ, Couger MB, Eccles D, Li B, Lieber M, MacManes MD, Ott M, Orvis J,Pochet N, Strozzi F, Weeks N, Westerman R, William T, Dewey CN,Henschel R, Leduc RD, Friedman N, Regev A. 2013. De novo transcriptsequence reconstruction from RNA-seq using the Trinity platform forreference generation and analysis. Nat Protoc 8:1494–1512. https://doi.org/10.1038/nprot.2013.084.7. Marchler-Bauer A, Bo Y, Han L, He J, Lanczycki CJ, Lu S, Chitsaz F,Derbyshire MK, Geer RC, Gonzales NR, Gwadz M, Hurwitz DI, Lu F,Marchler GH, Song JS, Thanki N, Wang Z, Yamashita RA, Zhang D, ZhengC, Geer LY, Bryant SH. 2017. CDD/SPARCLE: functional classification ofproteins via subfamily domain architectures. Nucleic Acids Res 45:D200–D203. https://doi.org/10.1093/nar/gkw1129.8. Altschul SF, Wootton JC, Gertz EM, Agarwala R, Morgulis A, Schaffer AA,Yu YK. 2005. Protein database searches using compositionally adjustedsubstitution matrices. FEBS J 272:5101–5109. https://doi.org/10.1111/j.1742-4658.2005.04945.x.9. Shi M, Lin XD, Tian JH, Chen LJ, Chen X, Li CX, Qin XC, Li J, Cao JP, EdenJS, Buchmann J, Wang W, Xu J, Holmes EC, Zhang YZ. 2016. Redefiningthe invertebrate RNA virosphere. Nature 540:539–543. https://doi.org/10.1038/nature20167.10. Hoang DT, Chernomor O, von Haeseler A, Minh BQ, Vinh LS. 2018.UFBoot2: improving the ultrafast bootstrap approximation. Mol Biol Evol35:518–522. https://doi.org/10.1093/molbev/msx281.Microbiology Resource AnnouncementVolume 9 Issue 26 e00380-20 mra.asm.org 3

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Diabrotica undecimpunctata virus 2, a Novel Small RNA Virus Discovered from Southern Corn Rootworm, Diabrotica undecimpunctata howardi Barber (Coleoptera: Chrysomelidae)

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