First Sin Nombre virus (Orthohantavirus sinnombreense) genome sequences from the Northwestern United States

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Catherine Grady , Jennifer Horton , Cody J. Lauritsen , Stephen Fawcett , Samuel M. Goodfellow , Hanna N. Oltean , View ORCID Profile M. Pilar Fernandez , View ORCID Profile Stephanie N. Seifert doi: https://doi.org/10.1101/2025.09.10.675434 Grant Rickard 1 Paul G. Allen School for Global Health, Washington State University , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site Ricardo Rivero 1 Paul G. Allen School for Global Health, Washington State University , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Ricardo Rivero A. Catherine Grady 1 Paul G. Allen School for Global Health, Washington State University , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site Jennifer Horton 1 Paul G. Allen School for Global Health, Washington State University , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site Cody J. Lauritsen 1 Paul G. Allen School for Global Health, Washington State University , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site Stephen Fawcett 1 Paul G. Allen School for Global Health, Washington State University , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site Samuel M. Goodfellow 2 Department of Pathology and Laboratory Medicine, Children’s Hospital Los Angeles , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site Hanna N. Oltean 3 Zoonotic and Vector-borne Disease Program, Washington State Department of Health , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site M. Pilar Fernandez 1 Paul G. Allen School for Global Health, Washington State University , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for M. Pilar Fernandez Stephanie N. Seifert 1 Paul G. Allen School for Global Health, Washington State University , USA Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Stephanie N. Seifert For correspondence: stephanie.seifert{at}wsu.edu Abstract Full Text Info/History Metrics Preview PDF Abstract We report the first Sin Nombre virus (SNV) genome sequences from the Northwestern United States and the first SNV sequences recovered from voles. Analysis of samples collected from 189 individual rodents revealed high SNV prevalence in the region and evidence of viral reassortment, highlighting ongoing viral diversification in rodents. Background Orthohantavirus sinnombreense (Sin Nombre Virus, SNV) is a member of the genus Orthohantavirus (family Hantaviridae) and the primary cause of Hantavirus Pulmonary Syndrome (HPS) in North America. First identified during a 1993 outbreak in the Four Corners region of the United States, SNV is linked to severe respiratory disease and high mortality( 1 ). As of 2024, more than 850 HPS cases have been reported in the United States, with a 36% case-fatality rate( 2 , 3 ). SNV is primarily maintained by Peromyscus spp. (deer mice), widespread rodents that are frequently associated with agricultural and peridomestic settings. Human infection usually results from inhalation of aerosolized viral particles from contaminated excreta( 4 , 5 ), with zoonotic risk influenced by ecological factors affecting rodent density( 6 ). While SNV is commonly detected in deer mice, several reported detections in sympatric rodent taxa and broad geographic distributions suggest greater complexity in virus maintenance( 7 , 8 ). Viral genomic surveillance can provide insights on viral evolution and spread( 9 ), yet there are fewer than 100 full SNV genomes published and none from the Northwestern United States( 10 ). Here, we report the first detection and genome sequences of SNV in the Northwestern United States and in voles. The Study Between June 2023 and August 2023, we live-trapped rodents using Sherman traps at farms and natural areas in the Palouse region of eastern Washington and western Idaho ( Figure 1 ), a major agricultural hub dominated by wheat and canola fields in the inland Northwestern United States. Sampling was conducted over three consecutive nights, providing repeated fecal sampling with mark-recapture, and lethal sample collection including tissue sampling on the third night. We collected and tested samples from 189 individual rodents across agricultural and natural landscapes in the Palouse. Sherman live traps were deployed at 8 farms and 2 forested sites in three 100 m X 30 m grids at each location. All captured rodents were identified, body and weight measurements collected, and either marked with an ear tag and released or euthanized following AVMA guidelines as approved by the Institutional Animal Care and Use Committee of Washington State University. Samples were collected under Idaho scientific collections permits #36112 and JOE19, and Washington state scientific collections permit SEIFERT 23-122. Download figure Open in new tab Figure 1. Locations of successful unique rodent sampling grids by land type; farms in yellow and natural areas in blue. Basemap is Sentinel-2 cloudless, EOX IT Services GmbH (CC BY 4.0). State boundary is U.S. Census Bureau Cartographic Boundary files (5m, public domain). We collected fecal samples on all trapping nights along with serum, lung, and bladder tissue on the third and final trapping night at each site. As no fecal samples tested positive for SNV by RT-qPCR on mark-recapture days, we report only data from the final capture for each individual rodent sampled. Across all sites, we collected samples from 2 creeping voles ( Microtus oregoni ), 18 montane voles ( Microtus montanus ), 4 western meadow voles ( Microtus drummondii ), 2 house mice ( Mus musculus ), 153 western deer mice ( Peromyscus sonoriensis ), 1 yellow pine chipmunk ( Neotamias amoenus ), and 9 least chipmunks ( Neotamias minimus ). We used recombinant nucleocapsid (N) protein from Sin Nombre virus (strain SN77734; BEI Resources, Manassas, Virginia, USA; NR-9670) containing a C-terminal histidine tag to assess seroreactivity in rodent serum samples. Briefly, the N protein was immobilized on Maxisorp-coated plates (ThermoFisher Scientific, Waltham, Massachusetts, USA; Cat# 439454), after which rodent serum diluted at 1:100 was applied. N protein bound antibodies were detected with horseradish peroxidase-conjugated goat anti-rat secondary antibody (AbCAM, Eugene, Oregon, USA; Cat #ab97057). Positive and negative thresholds for each plate were determined as the mean optical density of naïve serum controls plus 3 SD. Total RNA was extracted from tissue samples and fecal samples using the Zymo Quick-RNA MagBead kit (Zymo Research, Irvine, California, USA; Cat #R2132) according to the manufacturer’s protocol for each sample type. Presence of viral RNA was detected by RT-qPCR as described in Williamson et al . 2021( 11 ) with an internal control targeting the Peromyscus maniculatus hypoxanthine-guanine phosphoribosyltransferase (HPRT1) gene with in-house designed forward primer (5’-CAAAGCCTAAGAGGAGAGTTCA-3’), reverse primer (5’-GATGGCCGCAGAACTAGAA-3’), and probe (5HEX/AGGAGTCCC/ZEN/ATTGATGTTGCCAGT/3IABkFQ). Unexpectedly, montane voles captured on farmlands showed the highest prevalence overall, with 50% seroprevalence and 22.2% RT-qPCR positive lung tissue samples ( Table 1 ). Western deer mice tested positive by serology and RT-qPCR with higher prevalence on farms compared to natural areas (29.3% vs 20.0% seropositive, 9.4% vs 7.6% RT-qPCR positive lung tissue samples) and were the only species with an RT-qPCR positive fecal sample and bladder sample, in both cases from adult males ( Table 1 ). View this table: View inline View popup Download powerpoint Table 1. Serology and RT-qPCR results for Sin Nombre virus in rodent species sampled from farms and natural areas in the Palouse region of eastern Washington and western Idaho. Numbers indicate positives/total tested, with prevalence (%) in parentheses. Taxa with RT-qPCR positive samples in bold. ND = not done. We modeled SNV lung positivity by RT-qPCR as a binary outcome using logistic regression with mean bias reduction (brglm2 package in R v4.1.3) for species with at least one RT-qPCR positive lung tissue sample. Predictors were land type (Forest or Farm), sex, age (Adult or Juvenile), and species ( Microtus montanus or Peromyscus sonoriensis ). Males had significantly higher odds of RT-qPCR positivity in the lung tissue samples than females (odds ratio [OR] = 6.27, confidence interval [CI] = 1.3 - 30.1), whereas land type, species, and age showed smaller effect sizes. Elevated SNV prevalence in male deer mice aligns with known SNV ecology, though the precise mechanism is unknown( 12 ) and warrants further study. We sequenced SNV using a tiled amplicon scheme proposed by Goodfellow et al .( 13 ) on the Oxford Nanopore platform. Reads were quality-controlled, primers trimmed, reads mapped, and the consensus sequence extracted with an in-house assembly pipeline ( https://github.com/viralemergence/SNVler ). We recovered sequence data for all three SNV genome segments from 10 individual rodents including, for the first time, two montane voles with segment completeness ranging from 24-100% and depth between 6.8X-476.1X (GenBank Accession numbers TBD). To address persistent M-segment dropouts, we designed flanking primers (MsegFor 5’-GCAGGTAGCTGATCTCAAG-3’, and MsegR 5’-CCAGTCCATGTAAGAGGTAC-3’) for amplification and sequencing, improving assemblies and informing future refinement of the primer set for the Northwest. These are the first SNV genomes from the Northwestern United States, filling a critical knowledge gap( 2 ). Segment-wise alignments curated from NCBI Virus were analyzed in BEAST v1.10.5 (strict clock, exponential coalescent) with uniform tip sampling for incomplete collection dates( 14 ). Estimated clock rates were 1.19×10 −4 , 1.203×10 −4 , and 1.223×10 −4 subs/site/year, with root-to-tip regression resulting in R-squared values of 0.197 (p-value: 4.16e-04) for the S-segment, 0.067 (p-value: 0.0282) for the M-segment, and 0.257 (p-value: 1.056e-04) for the L-segment, suggesting a weak temporal signal. This likely reflects geographic population structure, reinforcing the need for broader sequencing to temporally resolve the SNV phylogeny. All Palouse sequences formed a clade closest to genomes from Montana collected in 2008/2009 ( Figure 2 )( 11 ). Download figure Open in new tab Figure 2. Phylogenetic reconstruction of the Washington samples of Orthohantavirus sinnombreense (SNV) in the context of viral diversity in the United States for the (A) S-segment, (B) M-segment, and (C) L-segment. The clade grouping the Washington samples generated in this study is highlighted in blue; MIMO indicates sequences from Microtus montanus and PESO indicates sequences from Peromyscus sonoriensis . Discrete phylogeography based on the S-segment tree with the same MCC parameters( 15 ) inferred introduction into Washington from Montana circa 1915 (HPD95%: 1873 – 1982), followed by local diversification ( Figure 3A ). Bayesian Stochastic Search Variable Selection analysis( 15 ) shows low Bayes Factor support for Montana-Washington movement, suggesting unsampled intermediates and illustrating the need for continued genomic surveillance in the Northwestern United States. Download figure Open in new tab Figure 3. Evolutionary dynamics of SNV showing (A) phylogeographic reconstruction of movements between discrete states using Markov Jumps, with curve weight reflecting Bayes Factor support and (B) tanglegram showing topology changes across segment trees, where connecting lines link variants and major shifts indicate reassortment. Topological discordance across viral segments supports viral reassortment in the local rodent population ( Figure 3B ). Within the Palouse clade, the montane vole ( Microtus montanus , MIMO) derived variants shifted position across segment trees, alternately clustering with or diverging from the local western deer mouse-derived variants, further supporting reassortment and suggesting cross-species transmission between montane voles and western deer mice in the Northwestern United States ( Figure 3B ). Conclusions We report on the first SNV genome sequences recovered from the Northwestern United States. Our findings highlight viral reassortment among sympatric rodent hosts of SNV, underscoring the complexity of SNV evolution and maintenance in rodents. Understanding how multiple hosts contribute to these reassortment events will be crucial for clarifying transmission dynamics in wild rodents and anticipating risks of zoonotic emergence. Acknowledgements This publication was made possible by cooperative agreement CDC-RFA-FT-23-0069 from the CDC’s Center for Forecasting and Outbreak Analytics. SNS, RR, and JH received support from US National Science Foundation (NSF DBI 2515340). The contents are solely the responsibility of the authors and do not necessarily represent the official views of the National Science Foundation or the Centers for Disease Control and Prevention. We thank Dr. Steven Bradfute for helpful discussions and guidance on recovering full viral genome sequences and the many field technicians who supported collections in the summer of 2023. Funder Information Declared Centers for Disease Control and Prevention, https://ror.org/042twtr12 , CDC-RFA-FT-23-0069 U.S. National Science Foundation, https://ror.org/021nxhr62 , 2515340 References 1. ↵ Nichol ST , Spiropoulou CF , Morzunov S , Rollin PE , Ksiazek TG , Feldmann H , et al. Genetic identification of a hantavirus associated with an outbreak of acute respiratory illness . Science . 1993 Nov 5 ; 262 ( 5135 ): 914 – 7 . OpenUrl Abstract / FREE Full Text 2. ↵ Whitmer SLM , Whitesell A , Mobley M , Talundzic E , Shedroff E , Cossaboom CM , et al. Human Orthohantavirus disease prevalence and genotype distribution in the U.S., 2008–2020: a retrospective observational study . Lancet Reg Health – Am [Internet] . 2024 Sep 1 [cited 2025 May 19]; 37 . Available from: https://www.thelancet.com/journals/lanam/article/PIIS2667-193X(24)00163-7/fulltext 3. ↵ Schmaljohn C , Hjelle B. Hantaviruses: a global disease problem . Emerg Infect Dis . 1997 Jun ; 3 ( 2 ): 95 – 104 . OpenUrl CrossRef PubMed Web of Science 4. ↵ Zeitz PS , Butler JC , Cheek JE , Samuel MC , Childs JE , Shands LA , et al. A case-control study of hantavirus pulmonary syndrome during an outbreak in the southwestern United States . J Infect Dis . 1995 Apr ; 171 ( 4 ): 864 – 70 . OpenUrl CrossRef PubMed Web of Science 5. ↵ Childs JE , Ksiazek TG , Spiropoulou CF , Krebs JW , Morzunov S , Maupin GO , et al. Serologic and genetic identification of Peromyscus maniculatus as the primary rodent reservoir for a new hantavirus in the southwestern United States . J Infect Dis . 1994 Jun ; 169 ( 6 ): 1271 – 80 . OpenUrl CrossRef PubMed Web of Science 6. ↵ Glass GE , Cheek JE , Patz JA , Shields TM , Doyle TJ , Thoroughman DA , et al. Using remotely sensed data to identify areas at risk for hantavirus pulmonary syndrome . Emerg Infect Dis . 2000 Jun ; 6 ( 3 ): 238 – 47 . OpenUrl CrossRef PubMed Web of Science 7. ↵ Goodfellow SM , Nofchissey RA , Schwalm KC , Cook JA , Dunnum JL , Guo Y , et al. Tracing Transmission of Sin Nombre Virus and Discovery of Infection in Multiple Rodent Species . J Virol . 2021 Nov 9 ; 95 ( 23 ): e01534 – 21 . OpenUrl PubMed 8. ↵ Goodfellow SM , Nofchissey RA , Arsnoe D , Ye C , Lee S , Park J , et al. Case of Human Orthohantavirus Infection, Michigan, USA, 2021 . Emerg Infect Dis . 2024 Apr ; 30 ( 4 ): 817 – 21 . OpenUrl PubMed 9. ↵ Hadfield J , Megill C , Bell SM , Huddleston J , Potter B , Callender C , et al. Nextstrain: real-time tracking of pathogen evolution . Bioinformatics . 2018 Dec 1 ; 34 ( 23 ): 4121 – 3 . OpenUrl CrossRef PubMed 10. ↵ Whitmer SLM , Whitesell A , Mobley M , Talundzic E , Shedroff E , Cossaboom CM , et al. Human Orthohantavirus disease prevalence and genotype distribution in the U.S., 2008–2020: a retrospective observational study . Lancet Reg Health – Am [Internet] . 2024 Sep 1 [cited 2024 Dec 16]; 37 . Available from: https://www.thelancet.com/journals/lanam/article/PIIS2667-193X(24)00163-7/fulltext 11. ↵ Williamson BN , Meade-White K , Boardman K , Schulz JE , Telford CT , Figueroa Acosta DM , et al. Continuing Orthohantavirus Circulation in Deer Mice in Western Montana . Viruses . 2021 Jun ; 13 ( 6 ): 1006 . OpenUrl PubMed 12. ↵ Warner BM , Stein DR , Griffin BD , Tierney K , Leung A , Sloan A , et al. Development and Characterization of a Sin Nombre Virus Transmission Model in Peromyscus maniculatus . Viruses . 2019 Feb ; 11 ( 2 ): 183 . OpenUrl PubMed 13. ↵ Goodfellow SM , Nofchissey RA , Ye C , Dunnum JL , Cook JA , Bradfute SB . Use of a Novel Detection Tool to Survey Orthohantaviruses in Wild-Caught Rodent Populations . Viruses . 2022 Mar 25 ; 14 ( 4 ). 14. ↵ Suchard MA , Lemey P , Baele G , Ayres DL , Drummond AJ , Rambaut A. Bayesian phylogenetic and phylodynamic data integration using BEAST 1.10 . Virus Evol [Internet] . 2018 Jan 1 [cited 2021 Jul 11]; 4 ( 1 ). Available from: https://academic.oup.com/ve/article/4/1/vey016/5035211 15. ↵ Lemey P , Rambaut A , Drummond AJ , Suchard MA . Bayesian Phylogeography Finds Its Roots . PLOS Comput Biol . 2009 Sep ; 5 ( 9 ): e1000520 . OpenUrl CrossRef PubMed View the discussion thread. Back to top Previous Next Posted September 16, 2025. Download PDF Email Thank you for your interest in spreading the word about bioRxiv. NOTE: Your email address is requested solely to identify you as the sender of this article. Your Email * Your Name * Send To * Enter multiple addresses on separate lines or separate them with commas. You are going to email the following First Sin Nombre virus (Orthohantavirus sinnombreense) genome sequences from the Northwestern United States Message Subject (Your Name) has forwarded a page to you from bioRxiv Message Body (Your Name) thought you would like to see this page from the bioRxiv website. 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