{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T02:10:00Z","timestamp":1790302200514,"version":"4.1.0"},"reference-count":70,"publisher":"Public Library of Science (PLoS)","issue":"4","license":[{"start":{"date-parts":[[2020,4,30]],"date-time":"2020-04-30T00:00:00Z","timestamp":1588204800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"DOI":"10.1371\/journal.pone.0231256","type":"journal-article","created":{"date-parts":[[2020,4,30]],"date-time":"2020-04-30T17:32:09Z","timestamp":1588267929000},"page":"e0231256","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":21,"title":["Modeling the Justinianic Plague: Comparing hypothesized transmission routes"],"prefix":"10.1371","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3852-5374","authenticated-orcid":true,"given":"Lauren A.","family":"White","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lee","family":"Mordechai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2020,4,30]]},"reference":[{"key":"pone.0231256.ref001","first-page":"48","volume-title":"Biology of man in history","author":"J-N Biraben","year":"1975"},{"key":"pone.0231256.ref002","doi-asserted-by":"crossref","DOI":"10.4324\/9781315255439","volume-title":"Famine and pestilence in the late Roman and early Byzantine Empire: a systematic survey of subsistence crises and epidemics","author":"DC Stathakopoulos","year":"2017"},{"key":"pone.0231256.ref003","doi-asserted-by":"crossref","DOI":"10.2307\/j.ctv9b2txr","volume-title":"The fate of Rome","author":"K. Harper","year":"2017"},{"key":"pone.0231256.ref004","doi-asserted-by":"crossref","first-page":"e1003349","DOI":"10.1371\/journal.ppat.1003349","article-title":"Yersinia pestis DNA from skeletal remains from the 6(th) century AD reveals insights into Justinianic Plague","volume":"9","author":"M Harbeck","year":"2013","journal-title":"PLoS Pathog"},{"key":"pone.0231256.ref005","first-page":"1","article-title":"Yersinia pestis and the plague of Justinian 541\u2013543 AD: A genomic analysis","volume":"3099","author":"DM Wagner","year":"2014","journal-title":"Lancet Infect Dis"},{"key":"pone.0231256.ref006","doi-asserted-by":"crossref","first-page":"2911","DOI":"10.1093\/molbev\/msw170","article-title":"A high-coverage Yersinia pestis genome from a sixth-century Justinianic Plague victim","volume":"33","author":"M Feldman","year":"2016","journal-title":"Mol Biol Evol"},{"key":"pone.0231256.ref007","doi-asserted-by":"crossref","first-page":"12363","DOI":"10.1073\/pnas.1820447116","article-title":"Ancient Yersinia pestis genomes from across Western Europe reveal early diversification during the First Pandemic (541\u2013750)","volume":"116","author":"M Keller","year":"2019","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pone.0231256.ref008","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1111\/emed.12152","article-title":"The \u2018Justinianic Plague\u2019: The economic consequences of the pandemic in the eastern Roman empire and its cultural and religious effects","volume":"24","author":"M Meier","year":"2016","journal-title":"Early Mediev Eur"},{"key":"pone.0231256.ref009","first-page":"107","volume-title":"Hommes et richesses dans l\u2019Empire byzantin","author":"J. Durliat","year":"1989"},{"key":"pone.0231256.ref010","doi-asserted-by":"crossref","DOI":"10.1093\/acprof:oso\/9780199264490.001.0001","volume-title":"Framing the early Middle Ages","author":"C. Wickham","year":"2005"},{"key":"pone.0231256.ref011","doi-asserted-by":"crossref","DOI":"10.1007\/978-1-349-24765-3","volume-title":"The making of Orthodox Byzantium, 600\u20131025","author":"M. Whittow","year":"1996"},{"key":"pone.0231256.ref012","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1093\/pastj\/gtz009","article-title":"Rejecting catastrophe: The case of the Justinianic Plague*","volume":"244","author":"L Mordechai","year":"2019","journal-title":"Past Present"},{"key":"pone.0231256.ref013","doi-asserted-by":"crossref","first-page":"25546","DOI":"10.1073\/pnas.1903797116","article-title":"The Justinianic Plague: An inconsequential pandemic?","volume":"116","author":"L Mordechai","year":"2019","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pone.0231256.ref014","doi-asserted-by":"crossref","first-page":"20160618","DOI":"10.1098\/rspb.2016.0618","article-title":"Epidemiological analysis of the Eyam plague outbreak of 1665\u20131666","volume":"283","author":"LK Whittles","year":"2016","journal-title":"Proc R Soc B Biol Sci"},{"key":"pone.0231256.ref015","doi-asserted-by":"crossref","first-page":"181695","DOI":"10.1098\/rsos.181695","article-title":"Epidemiology of a bubonic plague outbreak in Glasgow, Scotland in 1900","volume":"6","author":"KR Dean","year":"2019","journal-title":"R Soc Open Sci"},{"key":"pone.0231256.ref016","first-page":"201715640","article-title":"Human ectoparasites and the spread of plague in Europe during the Second Pandemic","volume":"115","author":"KR Dean","year":"2018","journal-title":"Proc Natl Acad Sci"},{"key":"pone.0231256.ref017","article-title":"The user\u2019s guide to comparative genomics with EnteroBase. Three case studies: Micro-clades within Salmonella enterica serovar Agama, ancient and modern populations of Yersinia pestis, and core genomic diversity of all Escherichia","author":"Z Zhou","year":"2019","journal-title":"bioRxiv"},{"key":"pone.0231256.ref018","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1086\/428617","article-title":"Risk of person-to-person transmission of pneumonic plague","volume":"40","author":"JL Kool","year":"2005","journal-title":"Clin Infect Dis"},{"key":"pone.0231256.ref019","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/S0140-6736(99)05163-6","article-title":"Epidemiological and diagnostic aspects of the outbreak of pneumonic plague in Madagascar","volume":"355","author":"M Ratsitorahina","year":"2000","journal-title":"Lancet"},{"key":"pone.0231256.ref020","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1162\/002219503322645439","article-title":"Rats, communications, and plague: Toward an ecological history","volume":"34","author":"M McCormick","year":"2003","journal-title":"J Interdiscip Hist"},{"key":"pone.0231256.ref021","doi-asserted-by":"crossref","first-page":"1752","DOI":"10.1016\/j.jas.2012.12.007","article-title":"Rats cannot have been intermediate hosts for Yersinia pestis during medieval plague epidemics in Northern Europe","volume":"40","author":"AK Hufthammer","year":"2013","journal-title":"J Archaeol Sci"},{"key":"pone.0231256.ref022","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1017\/CCOL0521817463.003","volume-title":"The Cambridge companion to the Age of Justinian","author":"B. Croke","year":"2005"},{"key":"pone.0231256.ref023","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1017\/CHOL9780521263351.028","volume-title":"The Cambridge ancient history XI: The high empire","author":"BW Frier","year":"2000"},{"key":"pone.0231256.ref024","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1017\/CCOL0521817463.006","volume-title":"The Cambridge companion to the Age of Justinian","author":"P. Horden","year":"2005"},{"key":"pone.0231256.ref025","doi-asserted-by":"crossref","first-page":"903","DOI":"10.1038\/35038073","article-title":"Metapopulation dynamics of bubonic plague","volume":"407","author":"MJ Keeling","year":"2000","journal-title":"Nature"},{"key":"pone.0231256.ref026","doi-asserted-by":"crossref","first-page":"2219","DOI":"10.1098\/rspb.2000.1272","article-title":"Bubonic plague: a metapopulation model of a zoonosis","volume":"267","author":"MJ Keeling","year":"2000","journal-title":"Proc R Soc London B Biol Sci"},{"key":"pone.0231256.ref027","doi-asserted-by":"crossref","first-page":"608","DOI":"10.3201\/eid1004.030509","article-title":"Epidemiologic determinants for modeling pneumonic plague outbreaks","volume":"10","author":"R Gani","year":"2004","journal-title":"Emerg Infect Dis"},{"key":"pone.0231256.ref028","doi-asserted-by":"crossref","first-page":"460","DOI":"10.3201\/eid1203.051051","article-title":"Pneumonic plague cluster, Uganda, 2004","volume":"12","author":"EM Begier","year":"2006","journal-title":"Emerg Infect Dis"},{"key":"pone.0231256.ref029","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/1756-3305-4-191","article-title":"Effects of temperature on the transmission of Yersinia pestis by the flea, Xenopsylla cheopis, in the late phase period","volume":"4","author":"AM Schotthoefer","year":"2011","journal-title":"Parasit Vectors"},{"key":"pone.0231256.ref030","doi-asserted-by":"crossref","first-page":"6236","DOI":"10.1073\/pnas.0510090103","article-title":"Classic flea-borne transmission does not drive plague epizootics in prairie dogs","volume":"103","author":"CT Webb","year":"2006","journal-title":"Proc Natl Acad Sci"},{"key":"pone.0231256.ref031","doi-asserted-by":"crossref","first-page":"e1003039","DOI":"10.1371\/journal.pcbi.1003039","article-title":"Host resistance, population structure and the long-term persistence of bubonic plague: Contributions of a modelling approach in the Malagasy focus","volume":"9","author":"F Gascuel","year":"2013","journal-title":"PLOS Comput Biol"},{"key":"pone.0231256.ref032","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1007\/s10393-010-0312-3","article-title":"Susceptibility to Yersinia pestis experimental infection in wild Rattus rattus, reservoir of plague in Madagascar","volume":"7","author":"C Tollenaere","year":"2010","journal-title":"Ecohealth"},{"key":"pone.0231256.ref033","doi-asserted-by":"crossref","first-page":"906","DOI":"10.1644\/1545-1542(2001)082<0906:IOIPON>2.0.CO;2","article-title":"Influences of introduced plague on North American mammals: Implications from ecology of plague in Asia","volume":"82","author":"DE Biggins","year":"2001","journal-title":"J Mammology"},{"key":"pone.0231256.ref034","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1017\/S0022172400008500","article-title":"The respective influences of temperature and moisture upon the survival of the rat flea (Xenopsylla cheopis) away from its host","volume":"23","author":"AW Bacot","year":"1924","journal-title":"J Hyg"},{"key":"pone.0231256.ref035","unstructured":"Frequently Asked Questions | Plague | CDC. 2018. Available: https:\/\/www.cdc.gov\/plague\/faq\/index.htmlLB\u2014pLww"},{"key":"pone.0231256.ref036","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1016\/j.epidem.2018.05.001","article-title":"The 2017 plague outbreak in Madagascar: Data descriptions and epidemic modelling","volume":"25","author":"VK Nguyen","year":"2018","journal-title":"Epidemics"},{"key":"pone.0231256.ref037","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1007\/978-94-024-0890-4_11","article-title":"Plague: Clinics, diagnosis and treatment","author":"V V Nikiforov","year":"2016","journal-title":"Adv Exp Med Biol"},{"key":"pone.0231256.ref038","doi-asserted-by":"crossref","first-page":"16","DOI":"10.3201\/eid2101.140564","article-title":"Epidemiology of human plague in the United States, 1900\u20132012","volume":"21","author":"KJ Kugeler","year":"2015","journal-title":"Emerg Infect Dis"},{"key":"pone.0231256.ref039","doi-asserted-by":"crossref","first-page":"561","DOI":"10.1148\/radiology.139.3.7232721","article-title":"Radiographic manifestations of plaque in New Mexico, 1975\u20131980. A review of 42 proved cases","volume":"139","author":"DJ Alsofrom","year":"1981","journal-title":"Radiology"},{"key":"pone.0231256.ref040","doi-asserted-by":"crossref","first-page":"2281","DOI":"10.1001\/jama.283.17.2281","article-title":"Plague as a biological weapon","volume":"283","author":"T V Inglesby","year":"2000","journal-title":"JAMA"},{"key":"pone.0231256.ref041","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1016\/S0733-8627(02)00005-6","article-title":"Threats in bioterrorism. I: CDC category A agents","volume":"20","author":"RG Darling","year":"2002","journal-title":"Emerg Med Clin North Am"},{"key":"pone.0231256.ref042","author":"M Keeling","year":"2008"},{"key":"pone.0231256.ref043","doi-asserted-by":"crossref","first-page":"505","DOI":"10.1146\/annurev.ento.50.071803.130337","article-title":"Natural history of plague: perspectives from more than a century of research","volume":"50","author":"KL Gage","year":"2005","journal-title":"Annu Rev Entomol"},{"key":"pone.0231256.ref044","article-title":"Sanitation in India","author":"JA Turner","year":"1914","journal-title":"The Times of India"},{"key":"pone.0231256.ref045","doi-asserted-by":"crossref","first-page":"855","DOI":"10.2307\/4142","article-title":"Density dependence in host-parasitoid models","volume":"50","author":"RM May","year":"1981","journal-title":"J Anim Ecol"},{"key":"pone.0231256.ref046","first-page":"1","article-title":"Solving differential equations in R: Package deSolve","volume":"33","author":"SKP Setzer","year":"2010","journal-title":"J Stat Softw"},{"key":"pone.0231256.ref047","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1016\/j.jtbi.2008.04.011","article-title":"A methodology for performing global uncertainty and sensitivity analysis in systems biology","volume":"254","author":"S Marino","year":"2008","journal-title":"J Theor Biol"},{"key":"pone.0231256.ref048","doi-asserted-by":"crossref","first-page":"20121018","DOI":"10.1098\/rsif.2012.1018","article-title":"Sensitivity analysis of infectious disease models: Methods, advances and their application","volume":"10","author":"J Wu","year":"2013","journal-title":"J R Soc Interface"},{"key":"pone.0231256.ref049","doi-asserted-by":"crossref","first-page":"229","DOI":"10.2307\/1403510","article-title":"Sensitivity and uncertainty analysis of complex models of disease transmission: An HIV model, as an example","volume":"62","author":"SM Blower","year":"1994","journal-title":"Int Stat Rev"},{"key":"pone.0231256.ref050","article-title":"lhs: Latin hypercube samples","author":"R. Carnell","year":"2019"},{"key":"pone.0231256.ref051","article-title":"sensitivity: Global sensitivity analysis of model outputs","author":"B Iooss","year":"2019"},{"key":"pone.0231256.ref052","volume-title":"A history of the later Roman Empire, AD 284\u2013641","author":"S. Mitchell","year":"2014"},{"key":"pone.0231256.ref053","volume-title":"The chronicle of Zuqn\u012bn parts III and IV A.D. 488\u2013775","author":"Ps. Dionysius of Tel-Mahre","year":"1999"},{"key":"pone.0231256.ref054","volume-title":"Secret history","author":"Prokopios","year":"1914"},{"key":"pone.0231256.ref055","volume-title":"Wars","author":"Prokopios","year":"1914"},{"key":"pone.0231256.ref056","volume-title":"Evagrios 6.8. The ecclesiastical history of Evagrius Scholasticus","author":"Evagrios (Scholasticus)","year":"2000"},{"key":"pone.0231256.ref057","article-title":"Chronique de Michel le Syrien: patriarche jacobite d\u2019Antioche (1166\u20131199)","author":"Michael the Syrian","year":"1899","journal-title":"Ernest Leroux"},{"key":"pone.0231256.ref058","doi-asserted-by":"crossref","first-page":"20182429","DOI":"10.1098\/rspb.2018.2429","article-title":"The third plague pandemic in Europe","volume":"286","author":"B Bramanti","year":"2019","journal-title":"Proc Biol Sci"},{"key":"pone.0231256.ref059","first-page":"125","article-title":"La colonisation de l\u2019Europe par le rat noir (Rattus rattus)","volume":"13","author":"F Audoin-Rouzeau","year":"1994","journal-title":"Rev Pal\u00e9obiologie"},{"key":"pone.0231256.ref060","first-page":"382","volume-title":"Handbuch der S\u00e4ugetiere Europas","author":"K. Becker","year":"1978"},{"key":"pone.0231256.ref061","doi-asserted-by":"crossref","first-page":"1099","DOI":"10.1098\/rsif.2007.1339","article-title":"Analysis of the sensitivity properties of a model of vector-borne bubonic plague","volume":"5","author":"M Buzby","year":"2008","journal-title":"J R Soc Interface"},{"key":"pone.0231256.ref062","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1002\/evl3.111","article-title":"To block or not to block: The adaptive manipulation of plague transmission","volume":"3","author":"S Gandon","year":"2019","journal-title":"Evol Lett"},{"key":"pone.0231256.ref063","first-page":"85","article-title":"A review of plague persistence with special emphasis on fleas","volume":"46","author":"J Wimsatt","year":"2009","journal-title":"J Vector Borne Dis"},{"key":"pone.0231256.ref064","doi-asserted-by":"crossref","first-page":"408","DOI":"10.7589\/0090-3558-43.3.408","article-title":"Modeling plague persistence in host-vector communities in California","volume":"43","author":"JE Foley","year":"2007","journal-title":"J Wildl Dis"},{"key":"pone.0231256.ref065","doi-asserted-by":"crossref","first-page":"14601","DOI":"10.1073\/pnas.1604985113","article-title":"Climatic and evolutionary drivers of phase shifts in the plague epidemics of colonial India","volume":"113","author":"JA Lewnard","year":"2016","journal-title":"Proc Natl Acad Sci"},{"key":"pone.0231256.ref066","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1163\/22134522-12340068","article-title":"Mysterious and mortiferous clouds: The climate cooling and disease burden of Late Antiquity","volume":"12","author":"TP Newfield","year":"2018","journal-title":"Late Antiq Archaeol"},{"key":"pone.0231256.ref067","author":"AJ Coale","year":"1984"},{"key":"pone.0231256.ref068","doi-asserted-by":"crossref","first-page":"1511","DOI":"10.1007\/s11538-006-9174-9","article-title":"Epidemiological models with non-exponentially distributed disease stages and applications to disease control","volume":"69","author":"Z Feng","year":"2007","journal-title":"Bull Math Biol"},{"key":"pone.0231256.ref069","volume-title":"Part I: Report","author":"EA Gait","year":"1913"},{"key":"pone.0231256.ref070","doi-asserted-by":"crossref","first-page":"43467","DOI":"10.1038\/srep43467","article-title":"Network theory may explain the vulnerability of medieval human settlements to the Black Death pandemic","volume":"7","author":"JM G\u00f3mez","year":"2017","journal-title":"Sci Rep"}],"container-title":["PLOS ONE"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0231256","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,4,30]],"date-time":"2020-04-30T17:32:29Z","timestamp":1588267949000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0231256"}},"subtitle":[],"editor":[{"given":"David A.","family":"Larsen","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2020,4,30]]},"references-count":70,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2020,4,30]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0231256","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,4,30]]}}}