Digital Commons Network^™

Coproid Predicts The Source Of Coprolites And Paleofeces Using Microbiome Composition And Host Dna Content, Maxime Borry, Bryan Cordova, Angela Perri, Marsha Wibowo, Tanvi Prasad Honap, Jada Ko, Kate Britton, Linus Girdland-Flink, Robert C. Power, Ingelise Stuijts, Domingo C. Salazar-García, Courtney Hofman, Richard Hagan, Thérèse Samdapawindé Kagoné, Nicolas Meda, Helene Carabin, David Jacobson, Karl Reinhard, Cecil Lewis, Aleksandar Kostic, Choongwon Jeong, Alexander Herbig, Alexander Hübner, Christina Warinner

Karl Reinhard Publications

Shotgun metagenomics applied to archaeological feces (paleofeces) can bring new insights into the composition and functions of human and animal gut microbiota from the past. However, paleofeces often undergo physical distortions in archaeological sediments, making their source species difficult to identify on the basis of fecal morphology or microscopic features alone. Here we present a reproducible and scalable pipeline using both host and microbial DNA to infer the host source of fecal material. We apply this pipeline to newly sequenced archaeological specimens and show that we are able to distinguish morphologically similar human and canine paleofeces, as well as non-fecal …

Diet Analysis Reveals Pre-Historic Meals Among The Loma San Gabriel At La Cueva De Los Muertos Chiquitos, Rio Zape, Mexico (600–800 Ce), Elisa Pucu, Julia Russ, Karl Reinhard

Karl Reinhard Publications

Coprolites have been a source of study for archeologists due to several reasons: they not only provide information on the life and nutritional habits of ancient individuals but also on their health. In this paper, we processed 10 coprolites collected at La Cueva de Los Muertos Chiquitos (600–800 CE), Rio Zape, Mexico, with acetolysis solution for pollen analysis. The number of pollen grains/gram of each coprolite sample was quantified along with the macroscopic remains of these samples. The main food item ingested by the population was maize, followed by Agave. Squash blossoms were also part of their food source …

Pinworm Research In The Southwest Usa: Five Decades Of Methodological And Theoretical Development And The Epidemiological Approach, Morgana Camacho, Karl Reinhard

Karl Reinhard Publications

Pinworms infected Ancestral Pueblo populations since early periods of occupation on the Colorado Plateau. The high prevalence of pinworm found in these populations was correlated with the habitation style developments through time. However, in previous studies, Turkey Pen Cave, an early occupation site, and Salmon Ruins, a late occupation site, exhibited prevalences that were anomalously low, suggesting that these sites were outliers. Alternatively, it is possible that the previous quantification method was not successful in detecting the real prevalence and eggs per gram, which led to inexact interpretations. The aims of this study were to verify if previous pinworm prevalences …

Genetic And Functional Analyses Of Virulence Potential Of An Escherichia Coli O157:H7 Strain Isolated From Super-Shedder Cattle, Lin Teng, Shinyoung Lee, Dongjin Park, Kwangcheol Casey Jeong

Department of Food Science and Technology: Faculty Publications

Shiga toxin (Stx)-producing Escherichia coli (STEC) O157:H7 is an enteric pathogen that causes life-threatening disease in humans, with cattle being major natural reservoirs. A group of STEC O157:H7 with a dramatic combination of high virulence potentials and super-shedder bovine origin have been isolated. Here, an STEC O157:H7 isolate, JEONG-1266, was analyzed by comparative genomics, stx genotyping, and phenotypic analyses. The phylogenetic typing and whole-genome comparison consistently showed that JEONG-1266 is genetically close to EC4115 (one of 2006 Spinach outbreak isolates) and SS17 (an isolate from super-shedder cattle) strains, all of which belong to lineage I/II and Clade 8. Both lineage …

A Mouse Model Of Human Tlr4 D299g/T399i Snps Reveals Mechanisms Of Altered Lps And Pathogen Responses, Katharina Richard, Kurt H. Piepenbrink, Kari Ann Shirey, Archana Gopalakrishnan, Shreeram Nallar, Daniel J. Prantner, Darren J. Perkins, Wendy Lai, Alexandra Vik, Vladimir Y. Toshchakov, Chiguang Feng, Rachel Fanaroff, Andrei E. Medvedev, Jorge C.G. Blanco, Stefanie N. Vogel

Department of Food Science and Technology: Faculty Publications

Two cosegregating single-nucleotide polymorphisms (SNPs) in human TLR4, an A896G transition at SNP rs4986790 (D299G) and a C1196T transition at SNP rs4986791 (T399I), have been associated with LPS hyporesponsiveness and differential susceptibility to many infectious or inflammatory diseases. However, many studies failed to confirm these associations, and transfection experiments resulted in conflicting conclusions about the impact of these SNPs on TLR4 signaling. Using advanced protein modeling from crystallographic data of human and murine TLR4, we identified homologous substitutions of these SNPs in murine Tlr4, engineered a knock-in strain expressing the D298G and N397I TLR4 SNPs homozygously, and characterized in vivo …

The Trust Principles For Digital Repositories, Dawei Lin, Jonathan Crabtree, Ingrid Dillo, Robert R. Downs, Rorie Edmunds, David Giaretta, Marisa De Giusti, Hervé L'Hours, Wim Hugo, Reyna Jenkyns, Varsha Khodiyar, Maryann E. Martone, Mustapha Mokrane, Vivek Navale, Jonathan Petters, Barbara Sierman, Dina V. Sokolova, Martina Stockhause, John Westbrook

Copyright, Fair Use, Scholarly Communication, etc.

As information and communication technology has become pervasive in our society, we are increasingly dependent on both digital data and repositories that provide access to and enable the use of such resources. Repositories must earn the trust of the communities they intend to serve and demonstrate that they are reliable and capable of appropriately managing the data they hold.

Following a year-long public discussion and building on existing community consensus , several stakeholders, representing various segments of the digital repository community, have collaboratively developed and endorsed a set of guiding principles to demonstrate digital repository trustworthiness. Transparency, Responsibility, User focus, …

Adaptations Of Interferon Regulatory Factor 3 With Transition From Terrestrial To Aquatic Life, Monica Angeletti, Wan-Ling Nicole Hsu, Nashaat Majo, Hideaki Moriyama, Etsuko Moriyama, Luwen Zhang

School of Biological Sciences: Faculty Publications

Interferon regulatory factor 3 (IRF3) and IRF7 are closely related IRF members and the major factors for the induction of interferons, a key component in vertebrate innate immunity. However, there is limited knowledge regarding the evolution and adaptation of those IRFs to the environments. Two unique motifs in IRF3 and 7 were identified. One motif, GASSL, is highly conserved throughout the evolution of IRF3 and 7 and located in the signal response domain. Another motif, DPHK, is in the DNA-binding domain. The ancestral protein of IRF3 and 7 seemed to possess the DPHK motif. In the ray-finned fish lineage, while …

Critical Thinking, Perspective-Taking, And Cultural Humility In Global Refugee Health Studies: A Peer Review Of Teaching Benchmark Portfolio, Julie A. Tippens

UNL Faculty Course Portfolios

This portfolio examines the third iteration of a course I developed for the Department of Child, Youth and Family Studies: CYAF 493 – Global Case Studies in Refugee Health and Wellbeing. The primary goal of this course is for students to think critically about the social and structural determinants of refugee health and wellbeing. I have also integrated three core course values to guide instruction: critical thinking, perspective-taking, and cultural humility. Student learning and engagement was analyzed through four areas: (1) a baseline assessment of students’ perceptions of refugees, (2) weekly homework reflections, (3) a book review assignment, and (4) …

Immunization Strategies In Networks With Missing Data, Samuel F. Rosenblatt, Jeffrey A. Smith, G. Robin Gauthier, Laurent Hébert-Dufresne

Department of Sociology: Faculty Publications

Network-based intervention strategies can be effective and cost-efficient approaches to curtailing harmful contagions in myriad settings. As studied, these strategies are often impractical to implement, as they typically assume complete knowledge of the network structure, which is unusual in practice. In this paper, we investigate how different immunization strategies perform under realistic conditions—where the strategies are informed by partially- observed network data. Our results suggest that global immunization strategies, like degree immunization, are optimal in most cases; the exception is at very high levels of missing data, where stochastic strategies, like acquaintance immunization, begin to outstrip them in minimizing outbreaks. …

Competing Forces Of Withdrawal And Disease Avoidance In The Risk Networks Of People Who Inject Drugs, Elspeth Ready, Patrick Habecker, Roberto Abadie, Bilal Khan, Kirk Dombrowski

Department of Sociology: Faculty Publications

We analyze a network of needle-sharing ties among 117 people who inject drugs (PWID) in rural Puerto Rico, using exponential random graph modeling to examine whether network members engage in partner restriction to lower their risk of contracting HIV or hepatitis C (HCV), or in informed altruism to prevent others from contracting these infections. Although sharing of used syringes is a significant risk factor for transmission of these diseases among PWID, we find limited evidence for partner restriction or informed altruism in the network of reported needle-sharing ties. We find however that sharing of needles is strongly reciprocal, and individuals …

Immunization Strategies In Networks With Missing Data, Samuel Frederick Rosenblatt, Jeffrey A. Smith, G. Robin Gauthier, Laurent Hébert-Dufresne

Department of Sociology: Faculty Publications

Network-based intervention strategies can be effective and cost-efficient approaches to cur- tailing harmful contagions in myriad settings. As studied, these strategies are often impracti- cal to implement, as they typically assume complete knowledge of the network structure, which is unusual in practice. In this paper, we investigate how different immunization strategies perform under realistic conditions—where the strategies are informed by partially-observed network data. Our results suggest that global immunization strategies, like degree immunization, are optimal in most cases; the exception is at very high levels of missing data, where stochastic strategies, like acquaintance immunization, begin to outstrip them in mini- …

Winter Use Of A Rock Crevice By Northern Long-Eared Myotis (Myotis Septentrionalis) In Nebraska, Jeremy A. White, Patricia Freeman, Hans W. Otto, Cliff Lemen

School of Natural Resources: Faculty Publications

We used a combination of acoustic detectors, thermal imaging cameras, and mist netting to confirm the use of a rock crevice by northern long-eared myotis (Myotis septentrionalis) in winter in northeastern Nebraska. Bats roosted in a large crevice in a rock face along the Missouri River, and 5 individuals were captured emerging from the opening of the crevice in March 2017. Our study is the first to confirm the use of a rock-crevice hibernaculum by M. septentrionalis and adds to the growing literature on the winter use of rock crevices by hibernating bats in North America. As white-nose …

Connectivity Differences Between Gulf War Illness (Gwi) Phenotypes During A Test Of Attention, Tomas Clarke, Jessie Jamieson, Patrick Malone, Rakib U. Rayhan, Stuart Washington, John W. Vanmeter, James N. Baraniuk

Department of Mathematics: Faculty Publications

One quarter of veterans returning from the 1990–1991 Persian Gulf War have developed Gulf War Illness (GWI) with chronic pain, fatigue, cognitive and gastrointestinal dysfunction. Exertion leads to characteristic, delayed onset exacerbations that are not relieved by sleep. We have modeled exertional exhaustion by comparing magnetic resonance images from before and after submaximal exercise. One third of the 27 GWI participants had brain stem atrophy and developed postural tachycardia after exercise (START: Stress Test Activated Reversible Tachycardia). The remainder activated basal ganglia and anterior insulae during a cognitive task (STOPP: Stress Test Originated Phantom Perception). Here, the role of attention …

Association Of Serum Γ-Tocopherol Levels With Mortality: The Multiethnic Cohort Study, Weiwen Chai, Gertraud Maskarinec, Adrian A. Franke, Kristine R. Monroeuniversity Of Southern California, Song-Yi Park, Laurence N. Kolonel, Lynne R. Wilkens, Loïc Le Marchand, Robert V. Cooney

Department of Nutrition and Health Sciences: Faculty Publications

Background/Objectives—γ-Tocopherol has unique properties that protect against nitrogen oxide-mediated cellular damage. To elucidate the potential role of γ-tocopherol in the aging process, we examined the associations of serum γ-tocopherol levels with all-cause and cause-specific mortality.

Subjects/Methods—Among participants in the biorepository subcohort of the Multiethnic Cohort Study, pre-cancer diagnostic serum γ-tocopherol levels were measured in a subset of 3904 men and 4461 women. Of these, 22.7% of men and 13.5% of women died during a mean follow- up time of 9.6±2.6 years. Hazard ratios (HRs) and 95% confidence intervals (95%CIs) for mortality associated with γ-tocopherol were estimated by Cox proportional hazards …

Chloroviruses, James L. Van Etten, Irina V. Agarkova, David D. Dunigan

Nebraska Center for Virology: Faculty Publications

Chloroviruses are large dsDNA, plaque-forming viruses that infect certain chlorella-like green algae; the algae are normally mutualistic endosymbionts of protists and metazoans and are often referred to as zoochlorellae. The viruses are ubiquitous in inland aqueous environments throughout the world and occasionally single types reach titers of thousands of plaque-forming units per ml of native water. The viruses are icosahedral in shape with a spike structure located at one of the vertices. They contain an internal membrane that is required for infectivity. The viral genomes are 290 to 370 kb in size, which encode up to 16 tRNAs and 330 …

Migration To The Us Among Rural Puerto Ricans Who Inject Drugs: Influential Factors, Sources Of Support, And Challenges For Harm Reduction Interventions, Roberto Abadie, Patrick Habecker, C. Gelpi-Acosta, Kirk Dombrowski

Department of Sociology: Faculty Publications

Background: While PWID of Puerto Rican origin have been migrating to the US for decades, the range of factors influencing their migration to the US and the resources they draw on to do so are not well understood. This is particularly true for rural Puerto Rican PWID, and the present study is the first empirical research to document migration patterns among this population. The specificities of their migration raise important challenges that need to be documented in order to implement more effective harm reduction policies at home (Puerto Rico) and abroad (US).

Methods: This paper draws from data obtained employing …

A Model For The Prediction Of Antimicrobial Resistance In Escherichia Coli Based On A Comparative Evaluation Of Fatty Acid Profiles, Randal S. Stahl, Bledar Bisha, Sebabrata Mahapatra, Jeffrey C. Chandler

United States Department of Agriculture Wildlife Services: Staff Publications

Antimicrobial resistance is a threat to agricultural production and public health. In this proof-of-concept study, we investigated predicting antimicrobial sensitive/resistant (S/R) phenotypes and host sources of Escherichia coli (n = 128) based on differential fatty acid abundance. Myristic (14:0), pentadecanoic acid (15:0), palmitic (16:0), elaidic (18:19) and steric acid (18:0) were significantly different (α = 0.05) using a two-way ANOVA for predicting nalidixic acid, ciprofloxacin, aztreonam, cefatoxime, and ceftazidime S/R phenotypes. Additionally, analyses of palmitoleic (16:1), palmitic acid (16:0), methyl palmitate (i-17:0), and cis-9,10-methyleneoctadecanoic acid (19:0Δ) showed these markers were significantly different (α = 0.05) between isolates obtained from cattle …

The R Journal (December 2019) 11(2): Complete Issue, The R Foundation

The R Journal

Editorial, Michael J. Kane

Contributed Research Articles

Using Web Services to Work with Geodata in R, Jan-Philipp Kolb

orthoDr: Semiparametric Dimension Reduction via Orthogonality Constrained Optimization, Ruoqing Zhu, Jiyang Zhang, Ruilin Zhao, Peng Xu, Wenzhuo Zhou, and Xin Zhang

coxed: An R Package for Computing Duration-Based Quantities from the Cox Proportional Hazards Model, Jonathan Kropko and Jeffrey J. Harden

Modeling Regimes with Extremes: The Bayesdfa Package for Identifying and Forecasting Common Trends and Anomalies in Multivariate Time-Series Data, Eric J. Ward, Sean C. Anderson, Luis A. Damiano, Mary E. Hunsicker, and Michael A. Litzow

Fitting Tails by the Empirical Residual …

Small Ruminant Health Intervention Calendar In Ethiopia, Mesfin Mekonnen, Ayalew Assefa, Tesfalem Nane, Firdawok Ayele, Asrat Arke, Belay Elias, Barbara Wieland

Daugherty Water for Food Global Institute: Faculty Publications

Contents

1 Background................................................................................................... 3
2 Developing the health intervention calendar............................................................ 4
The treatment calendar ......................................................... 4 Applying the calendar..................................................................... 5 4 References................................................................... 8

Serological Survey And Pathogen Exposure Of Adult Female White-Tailed Deer In The Western Dakotas, Katherine L. Moratz, Bailey S. Gullikson, Eric S. Michel, Daniel M. Grove, Jonathan A. Jenks, William Jensen

The Prairie Naturalist

Establishing baseline values for pathogen exposure and nutritional indices is necessary to monitor population health. However, little is known about white-tailed deer (Odocoileus virginianus) pathogen exposure and nutritional condition in the Northern Great Plains. Our objective was to assess pathogen exposure and establish nutritional indices for female white-tailed deer in Dunn and Grant counties, North Dakota and Perkins County, South Dakota. During 2014, we collected blood serum from 150 adult female white-tailed deer. Pathogens with the highest antibody prevalence included West Nile Virus (WNV; 85%), epizootic hemorrhagic disease (48%), and malignant catarrhal fever (32%). Serum values for creatine …

Editorial, Michael J. Kane

The R Journal

On behalf of the editorial board, I am pleased to present Volume 11, Issue 2 of the R Journal and my first issue as the Editor in Chief. This year, both Colin Gillespie and Catherine Healey join the Editorial Board, and Norm Matloff will rotate out. The R Journal continues to see increases in impact and popularity and this year we plan on making advances to better serve the community and streamline the publishing process to meet the increase in submissions we have seen over the last few years.

The Idspatialstats R Package: Quantifying Spatial Dependence Of Infectious Disease Spread, John R. Giles, Henrik Salje, Justin Lessler

The R Journal

Spatial statistics for infectious diseases are important because the spatial and temporal scale over which transmission operates determine the dynamics of disease spread. Many methods for quantifying the distribution and clustering of spatial point patterns have been developed (e.g. K-function and pair correlation function) and are routinely applied to infectious disease case occurrence data. However, these methods do not explicitly account for overlapping chains of transmission and require knowledge of the underlying population distribution, which can be limiting when analyzing epidemic case occurrence data. Therefore, we developed two novel spatial statistics that account for these effects to estimate: 1) …

Lassa Virus Circulating In Liberia: A Retrospective Genomic Characterisation, Michael R. Wiley, Lawrence S. Fakoli, Andrew G. Letizia, Stephen R. Welch, Jason T. Ladner, Karla Prieto, Daniel Reyes, Nicole Espy, Joseph A. Chitty, Catherine B. Pratt, Nicholas Di Paola, Fahn Taweh, Desmond Williams, Jon Saindon, William G. Davis, Ketan Patel, Mitchell Holland, Daniel Negrón, Ute Ströher, Stuart T. Nichol, Shanmuga Sozhamannan, Pierre E. Rollin, John Dogba, Tolbert Nyenswah, Fatorma Bolay, César G. Albariño, Mosoka Fallah, Gustavo Palacios

University of Nebraska's National Strategic Research Institute: Publications

Background An alarming rise in reported Lassa fever cases continues in west Africa. Liberia has the largest reported per capita incidence of Lassa fever cases in the region, but genomic information on the circulating strains is scarce. The aim of this study was to substantially increase the available pool of data to help foster the generation of targeted diagnostics and therapeutics.

Methods Clinical serum samples collected from 17 positive Lassa fever cases originating from Liberia (16 cases) and Guinea (one case) within the past decade were processed at the Liberian Institute for Biomedical Research using a targeted-enrichment sequencing approach, producing …

Preface For Special Section On Archaeoparasitology: A Global Perspective On Ancient Parasites And Current Research Projects, Jong Yil Chai, Min Sea, Karl Reinhard, Dong Hoon Shin

Karl Reinhard Publications

Archaeoparasitological research has rapidly developed in recent years, entering a new stage of improved understanding of our ancestors’ parasitic infections across the world. This progress is based on a variety of recently developed research techniques. During a period of rapid innovation since 2016, there has been no much opportunity for parasitologists to gather and share in-depth views about this novel research field. This was remedied by the 14th International Congress of Parasitology (ICOPA) held in Korea (Aug. 19-24, 2018; EXCO, Daegu). For archaeoparasitologists, ICOPA provided a unique opportunity to gather and exchange research and ideas. The participation was unprecedented in …

Confusing A Pollen Grain With A Parasite Egg: An Appraisal Of “Paleoparasitological Evidence Of Pinworm (Enterobius Vermicularis) Infection In A Female Adolescent Residing In Ancient Tehran”, Morgana Camacho, Karl Reinhard

Karl Reinhard Publications

There is often the risk of confusing pollen grains with helminth eggs from archaeological sites. Thousands to millions of pollen grains can be recovered from archaeological burial sediments that represent past ritual, medication and environment. Some pollen grain types can be similar to parasite eggs. Such a confusion is represented by the diagnosis of enterobiasis in ancient Iran. The authors of this study confused a joint-pine (Ephedra spp.) pollen grain with a pinworm egg. This paper describes the specific Ephedra pollen morphology that can be confused with pinworm eggs.

Pinworm Infection At Salmon Ruins And Aztec Ruins: Relation To Pueblo Iii Regional Violence, Karl Reinhard, Morgana Camacho

Karl Reinhard Publications

The study of coprolites has been a theme of archaeology in the American Southwest. A feature of archaeoparasitology on the Colorado Plateau is the ubiquity of pinworm infection. As a crowd parasite, this ubiquity signals varying concentrations of populations. Our recent analysis of coprolite deposits from 2 sites revealed the highest prevalence of infection ever recorded for the region. For Salmon Ruins, the deposits date from AD 1140 to 1280. For Aztec Ruins, the samples can be dated by artifact association between AD 1182-1253. Both sites can be placed in the Ancestral Pueblo III occupation (AD 1100-1300), which included a …

Male Burmese Pythons Follow Female Scent Trails And Show Sex-Specific Behaviors, Shannon A. Richard, Eric A. Tillman, John S. Humphrey, Michael L. Avery, M. Rockwell Parker

United States Department of Agriculture Wildlife Services: Staff Publications

Animals communicate with potential mates using species-specific signals, and pheromones are powerful sexual signals that modify conspecific behavior to facilitate mate location. Among the vertebrates, snakes are especially adept in mate searching via chemical trailing, which is particularly relevant given that many snake species are invasive outside their native ranges. Chemical signals used in mate choice are, thus, potentially valuable tools for management of invasive snake species. The Burmese python (Python bivittatus) is an invasive snake in the Florida Everglades where it is negatively impacting native fauna. In this study, we sought to: (i) determine if males can follow conspecific …

Variation In Host Home Range Size Decreases Rabies Vaccination Effectiveness By Increasing The Spatial Spread Of Rabies Virus, Katherine M. Mcclure, Amy T. Gilbert, Richard B. Chipman, Erin E. Rees, Kim M. Pepin

United States Department of Agriculture Wildlife Services: Staff Publications

1. Animal movement influences the spatial spread of directly transmitted wildlife disease through host-host contact structure. Wildlife disease hosts vary in home range- associated foraging and social behaviours, which may increase the spread and intensity of disease outbreaks. The consequences of variation in host home range movement and space use on wildlife disease dynamics are poorly understood, but could help to predict disease spread and determine more effective disease management strategies.
2. We developed a spatially explicit individual-based model to examine the effect of spatiotemporal variation in host home range size on the spatial spread rate, persistence and incidence of rabies virus …

Food Habits Of Coyotes (Canis Latrans) In The Valles Caldera National Preserve, New Mexico, Suzanne J. Gifford, Eric M. Gese, Robert R. Parmenter

United States Department of Agriculture Wildlife Services: Staff Publications

Wildlife managers needed to understand coyote (Canis latrans) ecology in order to develop management plans on the nascent Valles Caldera National Preserve in northern New Mexico. Managers concerned about low elk (Cervus elaphus) recruitment had observed an increase in sightings of coyotes and observations of coyote predation on elk calves. Our objective was to identify and quantify coyote diet, and assess the temporal variation in coyote diet on the Valles Caldera National Preserve, particularly as related to elk calf consumption. We examined coyote food habits using 1,385 scats analyzed monthly from May 2005 to November 2008. …

Genome-Wide Variation In Potyviruses, Deepti Nigam, Katherine Latourrette, Pedro F.N. Souza, Hernan Garcia Ruiz

Nebraska Center for Virology: Faculty Publications

Potyviruses (family Potyviridae, genus Potyvirus) are the result of an initial radiation event that occurred 6,600 years ago. The genus currently consists of 167 species that infect monocots or dicots, including domesticated and wild plants. Potyviruses are transmitted in a non-persistent way by more than 200 species of aphids. As indicated by their wide host range, worldwide distribution, and diversity of their vectors, potyviruses have an outstanding capacity to adapt to new hosts and environments. However, factors that confer adaptability are poorly understood. Viral RNA-dependent RNA polymerases introduce nucleotide substitutions that generate genetic diversity. We hypothesized that selection imposed by …