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The Justinianic Plague: an interdisciplinary review

Published online by Cambridge University Press:  10 September 2019

Merle Eisenberg
Lee Mordechai
National Socio-Environmental Synthesis Center, University of


This article is a detailed critical review of all the major scholarly publications in the rapidly expanding field of the Justinianic Plague published from 2000 through 2018. It updates the article in this journal by Dionysios Stathakopoulos from 2000, while also providing a detailed appraisal of the state of the field across all disciplines, including: literary studies, archaeology, DNA evidence, climatology, and epidemiology. We also identify the current paradigm for the Justinianic Plague as well as survey possible avenues forward for the field in the future.1

Copyright © Centre for Byzantine, Ottoman and Modern Greek Studies, University of Birmingham, 2019 

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We would like to thank Tim Newfield and the two anonymous reviewers for commenting on an earlier version of this paper. Princeton's Climate Change and History Research Initiative provided an instrumental platform for the refinement and discussion of the ideas behind this paper. This survey is comprehensive through 2018. Due to publication schedules we were able to include only brief references to a few studies from 2019. We use the term ‘Justinianic Plague’ in this article as a shorthand to refer to all outbreaks of plague from c. 541–750 since that remains the most commonly used term to discuss the first pandemic.


2 Stathakopoulos, D., ‘The Justinianic Plague revisited’, Byzantine and Modern Greek Studies 24 (2000) 256–76CrossRefGoogle Scholar.

3 M. Eisenberg et al., ‘The environmental history of the Late Antique West’, Late Antique Archaeology (2018); L. McMahon and A. Sargent, ‘Environmental history of the Late Antique Eastern Mediterranean’, Late Antique Archaeology (2018); Newfield, T. and Labuhn, I., ‘Realizing consilience in studies of pre-instrumental climate and pre-laboratory disease’, The Journal of Interdisciplinary History 48, no. 2 (2017) 218–19Google Scholar.

4 The conference resulted from conversations at the American Academy in Rome two years earlier. See Little, L., ‘Preface’, in Little, L. (ed.) Plague and the End of Antiquity: The Pandemic of 541–750 (Cambridge 2007) xiiGoogle Scholar.

5 For the resulting book see Little (ed.), Plague and the End of Antiquity.

6 For both his catalogue and his epidemiology section, Stathakopoulos, D., Famine and Pestilence in the Late Roman and Early Byzantine Empire: A Systematic Survey of Subsistence Crises and Epidemics (Aldershot 2004) 124–34Google Scholar.

7 See the discussion in Stathakopoulos, Famine and Pestilence, 111 which he did not follow in his catalogue. For example, some entries for the first outbreak (e.g. #114–118, pp. 290–4) are based on circumstantial evidence and only one (#114) refers to buboes, but without a specific place.

8 For other catalogues of environmental events that include the primary source text in translation, see for example: Ambraseys, N., Earthquakes in the Mediterranean and Middle East: A Multidisciplinary Study of Seismicity up to 1900 (Cambridge 2009)CrossRefGoogle Scholar; Guidoboni, E., Catalogue of Ancient Earthquakes in the Mediterranean Area up to the 10th Century (Rome 1994)Google Scholar; Guidoboni, E. and Comastri, A., Catalogue of Earthquakes and Tsunamis in the Mediterranean Area from the 11th to the 15th Century (Rome 2005)Google Scholar. The Guidoboni catalogues also include primary sources in the original languages.

9 M. Kulikowski, ‘Plague in Spanish Late Antiquity’, in Little (ed.), Plague and the End of Antiquity, 150–70; A. Dooley, ‘The plague and its consequences in Ireland’, in L. Little (ed.), Plague and the End of Antiquity, 215–28.

10 J. Maddicott, ‘Plague in seventh-century England’, in L. Little (ed.), Plague and the End of Antiquity, 185–90.

11 Seger, T., ‘The plague of Justinian and other scourges: An analysis of the anomalies in the development of the iron age population in Finland’, Fornvännen 77 (1982) 184–96Google Scholar; Robin, C-J., ‘Guerre et épidémie dans les royaumes d'Arabie du Sud, d'après une inscription datée (IIe s. de l’ère chrétienne)’, Comptes rendus des séances de l'Académie des Inscriptions et Belles-Lettres 136, no. 1 (1992) 215–34Google Scholar.

12 This is not to say that 750 was arbitrarily selected, but the emphasis on such a round number as an anno mirabilis is tempting. Conrad, L. I., ‘Arabic plague chronologies and treatises: Social and historical factors in the formation of a literary genre’, Studia Islamica, no. 54 (1981) 53Google ScholarPubMed, notes the last outbreak in 749; Dols, M., ‘Plague in early Islamic history’, Journal of the American Oriental Society 94, no. 3 (1974) 380–81CrossRefGoogle Scholar, notes that it was severe under the Umayyads, but plague did seem to continue afterwards as well. See the more recent M. Morony, ‘“For whom does the writer write?”: The first bubonic plague pandemic according to Syriac sources’, in Little (ed.), Plague and the End of Antiquity, 67. Dating the end of the plague in the mid-eighth century goes back to the early twentieth century, but the precise year remains debatable. See: Stathakopoulos, ‘The Justinianic plague revisited’, 259, 264, 268 and passim.

13 Morony, “‘For whom does the writer write?’”, 68–9.

14 See, for example, the contributions of H. Kennedy, ‘Justinianic plague in Syria and the archaeological evidence’, in Little (ed.), Plague and the End of Antiquity, 87–98; Maddicott, ‘Plague in seventh-century England.’

15 Durliat, J., ‘La peste du VIe siècle: pour un nouvel examen des sources byzantines’, in Abadie-Reynal, C., Morrisson, C., and Lefort, J. (eds.), Hommes et richesses dans l'Empire byzantine (Paris 1989) 107–19Google Scholar.

16 Sarris, P., ‘The Justinianic plague: origins and effects’, Continuity and Change 17,2 (2002) 169182CrossRefGoogle Scholar; P. Sarris, ‘Bubonic plague in Byzantium: The evidence of non-literary sources’, in Little (ed.), Plague and the End of Antiquity, 119–32. Little appears unaware of this, although Sarris noted it in the first footnote in his 2007 chapter. See Little, ‘Preface’, xiii.

17 See, for example, Meier, M., ‘The “Justinianic plague”: The economic consequences of the pandemic in the Eastern Roman Empire and its cultural and religious effects’, Early Medieval Europe 24.3 (2016) 274, 280CrossRefGoogle Scholar, who describes it as ‘important and highly pertinent’.

18 See Little, ‘Preface’, xii–xiii.

19 M. McCormick, ‘Toward a molecular history of the Justinianic pandemic’, L. Little (ed.), Plague and the End of Antiquity, 290–312 [290]. McCormick's notion of continued discoveries in plague studies features in his recent thought as well. See his introduction of Science of the Human Past, SoHP: Johannes Krause 2/16/17, 2017, at 00:00:15, 00:03:45, 00:04:25, 00:04:35.

20 R. Sallares, ‘Ecology, evolution, and epidemiology of plague’, in Little (ed.), Plague and the End of Antiquity, 231–89.

21 Wickham, C., Framing the Early Middle Ages: Europe and the Mediterranean 400–800 (Oxford 2005)CrossRefGoogle Scholar; Ward-Perkins, B., The Fall of Rome and the End of Civilization (Oxford 2005)Google Scholar; Little, ‘Preface’, xi–xii.

22 Rosen, W., Justinian's Flea: Plague, Empire, and the Birth of Europe (Penguin 2007)Google Scholar. Notably, Antoniou and Sinakos’ attempt to connect the scientific background for plague, demographic decline, and the end of antiquity was published in a leading Byzantine history journal, Byzantinische Zeitschrift. Their effort, however, was little noticed. See Antoniou, I. and Sinakos, A., ‘The sixth-century plague, Its repeated appearance until 746 AD and the explosion of the Rabaul Volcano’, Byzantinische Zeitschrift 98.1 (2008) 14CrossRefGoogle Scholar.

23 Kulikowski, M., ‘Justinian's flea: Plague, empire, and the birth of Europe’, History: Reviews of New Books 35.4 (2007) 148Google Scholar, described it as being ‘a wasted opportunity’; A. Kaldellis, ‘Plague and the end of antiquity: The pandemic of 541–750’, and: Justinian's flea: Plague, empire, and the birth of Europe (Review)’, Journal of Late Antiquity 1.2 (2008) 385–8CrossRefGoogle Scholar, asserted that it ‘is full of errors, misunderstandings, and outdated notions’; Treadgold, W., Review of Justinian's Flea: Plague, Empire, and the Birth of Europe, by Rosen, William, The Historian 71.1 (2009) 172–3Google Scholar claimed that the book ‘is so inaccurate and poorly written as almost to give common sense a bad name’.

24 Dattwyler, R., ‘Book review’, New England Journal of Medicine 357, no. 13 (2007) 1354–5CrossRefGoogle Scholar; Burki, T., ‘Book: Justinian's flea: Plague, empire and the birth of Europe’, The Lancet Infectious Diseases; London 7.12 (2007)Google Scholar.

26 Roosen, J. and Curtis, D. R., ‘Dangers of non-critical use of historical plague data’, Emerging Infectious Diseases 24.1 (2018) 103–10CrossRefGoogle Scholar. And for the accessibility question: King, H. and Green., M. H.On the misuses of medical history’, The Lancet 391, no. 10128 (7 April 7 2018): 1354–5CrossRefGoogle ScholarPubMed.

27 On this point, see Stathakopoulos, D., ‘Death in the countryside: some thoughts on the effects of famine and epidemics’, Antiquité Tardive 20 (2012) 107CrossRefGoogle Scholar; in more detail: Bachrach, B., ‘Plague, population, and economy in Merovingian Gaul’, Journal of the Australian Early Medieval Association 3 (2007) 2957Google Scholar; Cameron, A., Procopius and the Sixth Century (Berkeley 1985) 3943Google Scholar.

28 The specific causal link between canons and plague, however, does not necessarily exist, since the vocabulary for plague never appears. See McCormick, M., ‘Tracking mass death during the fall of Rome's empire (I)’, Journal of Roman Archaeology 28 (2015) 338CrossRefGoogle Scholar, for one example. And see now: Gruber, H., ‘Indirect Evidence for the Social Impact of the Justinianic Pandemic: Episcopal Burial and Conciliar Legislation in Visigothic Hispania’, Journal of Late Antiquity 11 (2018), 193215CrossRefGoogle Scholar.

29 Conrad's works remain indispensable for the region, see: L. I. Conrad, ‘The plague in the early Medieval Near East’ (Princeton University 1981); Conrad, L. I, ‘Die Pest und ihr Soziales Umfeld im Nahen Osten des Frühen Mittelalters’, Der Islam 73.1 (1996) 81112CrossRefGoogle Scholar.

30 For the waves: Stathakopoulos, Famine and Pestilence, 110 his list is at 113–24; for the geographic range: 116.

31 Durliat, ‘La peste du VIe siècle’.

32 Sarris, ‘The Justinianic plague’, 173–8.

33 Sarris, ‘The Justinianic plague’, 169–73, 178–9.

34 Descriptions and discussions in Benovitz, N., ‘The Justinianic plague: evidence from the dated Greek epitaphs of Byzantine Palestine and Arabia’, Journal of Roman Archaeology 27 (2014) 491–2CrossRefGoogle Scholar; McCormick, ‘Tracking mass death (I)’, 327–8; Meier, ‘The “Justinianic plague”’, 267–9. The building inscription from Syria is understood to refer to a bubo (πóτμον βονβῶνος [sic] καὶ μάλης, which McCormick translates as ‘an evil destiny of bubo and armpit’); an epitaph from Spain mentions that the person died ab inguinali plaga (‘from the affliction of the groin’).

35 Benovitz, ‘The Justinianic plague’, 495–96. The number of dead for 541, the beginning of the first outbreak, is sixteen - double the number of dead in any other year in the studied interval between 300–699.

36 For two examples: Colt, H., Excavations at Nessana (Princeton 1962) 168 no. 80 and 179–81 nos. 112–14Google Scholar; Glucker, C., The City of Gaza in the Roman and Byzantine Periods (Oxford 1987) 124–7 nos. 9–11Google Scholar. Glucker's proof of plague is a citation to Colt, who simply made the suggestion, but left the question open.

37 McCormick, M., ‘Rats, communications, and plague: Toward an ecological history’, The Journal of Interdisciplinary History 34, no. 1 (2003) 125CrossRefGoogle Scholar; see also Stathakopoulos, D., ‘Invisible protagonists: The Justinianic plague from a zoocentric point of view’, in Anagnostakis, E., Kolias, T., and Papadopoulou, E. (eds.), Zώα και περιβάλλον στο Βυζάντιο (7ος-12ος αι.) = Animals and Environment in Byzantium (7th-12th c.), 87–95 (Athens 2011) 8795Google Scholar.

38 McCormick, ‘Rats, communications, and plague’, 9 knew of 65 finds of rat bones until the eighth century CE.

39 K. Royer, ‘The blind men and the elephant: Imperial medicine, medieval historians and the role of rats in the historiography of plague’, Medicine and Colonialism (2014) 113–24. Royer points out that even in India the Third Pandemic varied in its epidemiology and notes that historians prefer to seek a one-size-fits-all explanation for complex and protean biological phenomena. For a recent study about the Black Death, see: Dean, K. et al. , ‘Human ectoparasites and the spread of plague in Europe during the Second Pandemic’, Proceedings of the National Academy of Sciences 115.6 (2018) 1304–9CrossRefGoogle ScholarPubMed.

40 Dean, K., Krauer, F., and Schmid, B., ‘Epidemiology of a bubonic plague outbreak in Glasgow, Scotland in 1900’, Royal Society Open Science 6.1 (2019) 3CrossRefGoogle ScholarPubMed.

41 Sallares, ‘Ecology, evolution, and epidemiology of plague’, 269. This work does not appear to be comprehensive and Sallares maintained his support for the rat-and-flea model.

42 Cohn, S., ‘The Black Death: εnd of a paradigm’, The American Historical Review 107.3 (2002) 703–38CrossRefGoogle Scholar; Cohn, S., ‘Epidemiology of the Black Death and successive waves of plague’, Medical History. Supplement, no. 27 (2008) 7577Google Scholar, also discusses the discrepancies between the Justinianic Plague and the Third Pandemic; see also Horden, P., ‘Mediterranean plague in the age of Justinian’, in Maas, M. (ed.), The Cambridge Companion to the Age of Justinian (Cambridge 2005) 134–60CrossRefGoogle Scholar; Royer, ‘The blind men and the elephant’.

43 While Stathakopoulos, Famine and Pestilence, 112 suggests that the next two pandemics should be studied, he cited nine recent works on the Black Death, but only one (from 1936) on the Third Pandemic.

44 Kool, J., ‘Risk of person-to-person transmission of pneumonic plague’, Clinical Infectious Diseases: An Official Publication of the Infectious Diseases Society of America 40.8 (2005) 1166–72CrossRefGoogle ScholarPubMed. For an argument against pneumonic, see Begier, E. et al. , ‘Pneumonic plague cluster, Uganda, 2004’, Emerging Infectious Diseases 12.3 (2006) 460–67CrossRefGoogle Scholar.

45 See, for example, Drancourt, M., Houhamdi, L., and Raoult, D., ‘Yersinia Pestis as a telluric, human ectoparasite-borne organism’, The Lancet Infectious Diseases 6, no. 4 (2006) 234–41CrossRefGoogle ScholarPubMed; Hufthammer, A. and Walløe, L., ‘Rats cannot have been intermediate hosts for Yersinia Pestis during medieval plague epidemics in Northern Europe’, Journal of Archaeological Science 40, no. 4 (2013) 1752–59CrossRefGoogle Scholar; Houhamdi, L. and Raoult, D., ‘Different genes govern Yersinia Pestis pathogenicity in Caenorhabditis Elegans and human lice’, Microbial Pathogenesis 44.5 (2008) 435–37CrossRefGoogle ScholarPubMed; Piarroux, R. et al. , ‘Plague epidemics and lice, Democratic Republic of the Congo’, Emerging Infectious Diseases 19.3 (2013) 505–6CrossRefGoogle ScholarPubMed.

46 Cohn, ‘Epidemiology’, 95–7; Campbell, B. M. S., The Great Transition (Cambridge 2016) 232–3CrossRefGoogle Scholar.

47 See for example K. Dean, ‘Modeling plague transmission in medieval European cities’ (University of Oslo, 2015),; Dean et al., ‘Human ectoparasites’.

48 Although historians were well aware that they could not resolve this question by themselves, they made arguments for both places of origin. Tsiamis, C., Poulakou-Rebelakou, E. and Petridou, E., ‘The Red Sea and the port of Clysma. A possible gate of Justinian's Plague’, Gesnerus 66.2 (2009) 209–17Google ScholarPubMed, argued that plague reached Pelusium through the port of Clysma in the Red Sea; For an African origin, see Sarris, ‘Bubonic Plague in Byzantium’, 120–3. For a central Asian origin, see for example: Green, M. et al. , ‘Yersinia Pestis and the three plague pandemics’, The Lancet Infectious Diseases; London 14.10 (2014) 918CrossRefGoogle ScholarPubMed; Kutyrev, V. et al. , ‘Phylogeny and classification of Yersinia Pestis through the lens of strains from the plague foci of Commonwealth of Independent States’, Frontiers in Microbiology 9 (2018) 1106CrossRefGoogle ScholarPubMed.

49 Cui, Y. et al. , ‘Historical variations in mutation rate in an epidemic pathogen, Yersinia Pestis’, Proceedings of the National Academy of Sciences of the United States of America 110.2 (2013) 577–82CrossRefGoogle Scholar. The conclusion has been accepted by historians, e.g. Green, M., ‘Taking “pandemic” seriously: making the Black Death global,’ The Medieval Globe 1 (2014) 36–7Google Scholar; Varlık, N., Plague and Empire in the Early Modern Mediterranean World: The Ottoman Experience, 1347–1600 (Cambridge 2015) 1920CrossRefGoogle Scholar (emphasizing the origins of rats); Harper, K., The Fate of Rome: Climate, Disease, and the End of an Empire (Princeton 2017), 210–14CrossRefGoogle Scholar.

50 Such questions have been asked, for instance by: Horden, ‘Mediterranean plague’; J. Hays, ‘Historians and epidemics: Simple questions, complex answers’, in Little (ed.), Plague and the End of Antiquity, 42ff.

51 See, for example, the hints in Horden, ‘Mediterranean plague’, 143ff and his stronger conclusion that the Justinianic Plague ‘absorbs a great deal of our energy; it gives very little out’ (p. 157).

52 For the positive DNA tests, see Drancourt, M. et al. , ‘Yersinia Pestis Orientalis in remains of ancient plague patients’, Emerging Infectious Diseases 13, no. 2 (2007) 332–3CrossRefGoogle ScholarPubMed; and for the archaeological evidence: Signoli, M. et al. , ‘Une sépulture de pestiférés du Haut Moyen Âge à Vienne (Isère)’, Archéologie du Midi médiéval 27.1 (2009) 1929CrossRefGoogle Scholar. See now also Keller, et al. ‘Ancient Yersinia pestis genomes from across Western Europe reveal early diversification during the First Pandemic (541–750)’, Proceedings of the National Academy of Sciences, 116.25 (2019) 1236312372CrossRefGoogle Scholar.

53 McCormick, ‘Tracking mass death (I)’; McCormick, M., ‘Tracking mass death during the fall of Rome's empire (II): A first inventory of mass graves’, Journal of Roman Archaeology 29 (2016) 1004–7CrossRefGoogle Scholar, for the catalogue of mass graves.

54 McCormick, ‘Tracking mass death (I)’, 333.

55 Gutsmiedl-Schümann, D. et al. , ‘Digging up the plague: A diachronic comparison of aDNA confirmed plague burials and associated burial customs in Germany’, Praehistorische Zeitschrift 92.2 (2018), 408, 410, 416CrossRefGoogle Scholar; Ivison, E., ‘Burial and urbanism at Late Antique and early Byzantine Corinth (c. AD 400–700)’, in Christie, N. and Loseby, S. T. (eds.), Towns in Transition: Urban Evolution in Late Antiquity and the Early Middle Ages (Brookfield, VΤ 1996), 108–10, 112–14Google Scholar; Schneider, T., ‘Mehrfachbestattungen von Männern in der Merowingerzeit’, Zeitschrift für Archäologie des Mittelalters 36 (2008) 132Google Scholar, for Merovingian graves in particular.

56 Wickham, Framing the Early Middle Ages; McCormick, M., Origins of the European Economy: Communications and Commerce, A.D. 300–900 (Cambridge 2001)Google Scholar.

57 Kennedy, ‘Justinianic Plague in Syria’.

58 Avni, G., The Byzantine-Islamic Transition in Palestine: An Archaeological Approach (Oxford 2014)CrossRefGoogle Scholar; Magness, J., The Archaeology of the Early Islamic Settlement in Palestine (Winona Lake, IN 2003)Google Scholar.

59 The connection between Y. pestis and the historical pandemics was assumed until called into question in 1984. For a review of the debate, see Little, L., ‘Plague historians in lab coats’, Past & Present 213.1 (2011): 272ffCrossRefGoogle ScholarPubMed. For a useful review of scientific research on plague, see Gage, K. and Kosoy, M., ‘Natural history of plague: Perspectives from more than a century of research’, Annual Review of Entomology 50 (2005) 505–28CrossRefGoogle Scholar. See below for a survey of ancient DNA, which is comprehensive through August 2018, but only discusses a few key updates afterward.

60 Drancourt, M. et al. , ‘Genotyping, Orientalis-like Yersinia Pestis, and plague pandemics’, Emerging Infectious Diseases 10, no. 9 (2004) 1585–92CrossRefGoogle ScholarPubMed; for Vienne: Drancourt et al., ‘Yersinia Pestis Orientalis’. For Poitiers, see: Castex, D. and Kacki, S., ‘Demographic patterns distinctive of epidemic cemeteries in archaeological samples’, Microbiology Spectrum 4, no. 4 (2016)Google ScholarPubMed, which is based on material culture remains and not DNA.

61 For problems: Vergnaud, G., ‘Yersinia Pestis genotyping’, Emerging Infectious Diseases 11.8 (2005) 1317–19Google ScholarPubMed; Prentice, M., Gilbert, T., and Cooper, A., ‘Was the Black Death caused by Yersinia Pestis?’, The Lancet. Infectious Diseases 4.2 (2004) 72CrossRefGoogle ScholarPubMed; and most notably: Gilbert, M. T. P. et al. , ‘Absence of Yersinia Pestis- specific DNA in human teeth from five European excavations of putative plague victims’, Microbiology 150.2 (2004) 341–54CrossRefGoogle ScholarPubMed. Harbeck, M. et al. , ‘Yersinia Pestis DNA from skeletal remains from the 6th century AD reveals insights into Justinianic Plague’, PLOS Pathogens 9.5 (2013)CrossRefGoogle Scholar; further critiqued Drancourt et al.’s 2007 paper. On more recent French work, see the special journal issue edited by: Drancourt, M. and Raoult, D. (eds.), Paleomicrobiology of Humans (Washington, D.C. 2016)CrossRefGoogle Scholar. For a personal discussion of these debates, see Raoult, D., ‘A personal view of how paleomicrobiology aids our understanding of the role of lice in plague pandemics’, Microbiology Spectrum 4.4 (2016)Google ScholarPubMed. On Italy and France from the High Middle Ages: Tran, T-N-N. et al. , ‘High throughput, multiplexed pathogen detection authenticates plague waves in medieval Venice, Italy’, PLoS ONE 6.3 (2011)CrossRefGoogle ScholarPubMed; Tran, T-N-N. et al. , ‘Brief communication: Co-detection of Bartonella Quintana and Yersinia Pestis in an 11th-15th burial site in Bondy, France’, American Journal of Physical Anthropology 145.3 (2011): 489–94CrossRefGoogle Scholar.

62 Wiechmann, I. and Grupe, G., ‘Detection of Yersinia pestis DNA in two early medieval skeletal finds from Aschheim (Upper Bavaria, 6th century A.D.)’, American Journal of Physical Anthropology 126.1 (2005): 4855CrossRefGoogle Scholar; Harbeck et al., ‘Yersinia Pestis DNA’; Wagner, D., ‘Yersinia Pestis and the plague of Justinian 541–543 AD: A genomic analysis’, The Lancet Infectious Diseases 14.4 (2014): 319–26CrossRefGoogle ScholarPubMed.

63 Feldman, M. et al. , ‘A high-coverage Yersinia Pestis genome from a sixth-century Justinianic Plague victim’, Molecular Biology and Evolution 33.11 (2016): 2911–23CrossRefGoogle ScholarPubMed.

64 Keller et al. ‘Ancient Yersinia pestis genomes from across Western Europe’, has the total numbers.

65 Gutsmiedl-Schümann et al., ‘Digging up the plague’. The quote is from p. 410 and refers to the Aschheim cemetery, although the same phenomenon is discussed for Altenerding as well.

66 Devignat, R., ‘Variétés de l'espèce Pasteurella Pestis’, Bulletin of the World Health Organization 4.2 (1951) 247–63Google Scholar.

67 For the most recent phylogenetic trees, see de Barros Damgaard, P. et al. , ‘137 ancient human genomes from across the Eurasian steppes’, Nature 557, no. 7705 (2018) fig. 9Google Scholar (Extended Data); and the synthesis with several descriptions of branches on the larger tree in Kutyrev et al., ‘Phylogeny and classification’. Another updated version is in pre-print: Zhou, et al., ‘The user's 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’, bioRxiv (2019). For discussions, see Green et al., ‘Yersinia Pestis and the three plague pandemics’; Sussman, G., ‘Scientists doing history: Central Africa and the origins of the First Plague Pandemic’, Journal of World History 26.2 (2016) 325–54CrossRefGoogle Scholar.

68 Büntgen, U. et al. , ‘Cooling and societal change during the Late Antique Little Ice Age from 536 to around 660 AD’, Nature Geoscience 9.3 (2016) 231–6CrossRefGoogle Scholar. For previous linkages between plague and climate, see Stenseth, N. et al. , ‘Plague dynamics are driven by climate variation’, Proceedings of the National Academy of Sciences of the United States of America 103.35 (2006) 13110–15CrossRefGoogle ScholarPubMed; Schmid, B. et al. , ‘Climate-driven introduction of the Black Death and successive plague re-introductions into Europe’, Proceedings of the National Academy of Sciences 112.10 (2015) 30203025CrossRefGoogle Scholar.

69 Harper, The Fate of Rome, 167–75, 218–20; K. Harper, ‘Invisible environmental history: Infectious disease in Late Antiquity’, Late Antique Archaeology, (2018). The catastrophist-determinist connection between the volcanic activity and the ensuing societal disruption and collapse world was discussed in a sensational form by Keys almost two decades ago. See Keys, D., Catastrophe: An Investigation into the Origins of the Modern World (London 1999)Google Scholar. For a more scholarly approach, which lacked some of the data now available, see: Arjava, A., ‘The mystery cloud of 536 CE in the Mediterranean sources’, Dumbarton Oaks Papers 59 (2005) 7394CrossRefGoogle Scholar.

70 On the difficulties in establishing such a connection, see: T. Newfield, ‘Mysterious and mortiferous clouds: The climate cooling and disease burden of Late Antiquity’, Late Antique Archaeology (2018).

71 Xu, L. et al. , ‘Wet climate and transportation routes accelerate the spread of human plague’, Proceedings of the Royal Society B: Biological Sciences 281, no. 1780 (2014) 19CrossRefGoogle ScholarPubMed; also, Snäll, T., Benestad, R. E., and Stenseth, N. C., ‘Expected future plague levels in a wildlife host under different scenarios of climate change’, Global Change Biology 15.2 (2009) 500–7CrossRefGoogle Scholar.

72 Stenseth et al., ‘Plague dynamics’ for warmer springs and wetter summers; Schmid et al., ‘Climate-driven introduction of the Black Death’ for cooling after a warm period.

73 Haldon, J. et al. , ‘The climate and environment of Byzantine Anatolia: Integrating science, history, and archaeology’, Journal of Interdisciplinary History 45.2 (2014) 113–61CrossRefGoogle Scholar; Izdebski, A., A Rural Economy in Transition: Asia Minor from Late Antiquity into the Early Middle Ages (Warsaw 2013)Google Scholar.

74 Xu, L. et al. , ‘Nonlinear effect of climate on plague during the Third Pandemic in China’, Proceedings of the National Academy of Sciences 108.25 (2011) 10214–19CrossRefGoogle Scholar; Xu et al., ‘Wet climate and transportation routes’.

75 On this, and for a broader nuanced discussion about the connection between plague and climate, see Newfield, ‘Mysterious and mortiferous clouds: The climate cooling and disease burden of Late Antiquity.’

76 Stathakopoulos, ‘The Justinianic Plague revisited’, 262ff.

77 Stathakopoulos, Famine and Pestilence, 139–41; repeated in Stathakopoulos, D., ‘Population, demography, and disease,’ in Jeffreys, E., Haldon, J. and Cormack, R. (eds.), The Oxford Handbook of Byzantine Studies (Oxford 2008) 310Google Scholar. An estimate found separately in Zuckerman, C., Du village à l'empire: autour du registre fiscal d'Aphroditô (525/526) (Paris 2004), 189212Google Scholar but who noted the population seems to have rebounded within a decade.

78 Horden, ‘Mediterranean plague’, 149; Antoniou and Sinakos, ‘The sixth-century plague.’

79 Kool, ‘Risk of person-to-person transmission’, 1166. The number is not cited in Kool's source for the paragraph.

80 Wagner, ‘Yersinia Pestis and the Plague of Justinian.’ Their estimate is based upon Procopius’ Secret History, which they admit ‘is disputed’.

81 Harper, The Fate of Rome, 244.

82 Mitchell, S., A History of the Later Roman Empire, AD 284–641, 2nd edn (Chichester 2015), 410–13, 479–91Google Scholar. Notably, Mitchell divides sites between pre-and post-550 dating to suggest that a demographic decline is a direct result of the plague. However, the imprecision in dating material culture through archaeological evidence, alongside the other potential causes for demographic decline in the mid-sixth century, cast doubt on these conclusions. Devoery, J-P., ‘Catastrophe, crise et changement social: à propos des paradigmes d'interprétation du développement médiéval (500–1100)’, in Buchet, L., Rigeade, C. and Séguy, I. (eds.), Actes des 9e Journées Anthropologiques de Valbonne (Valbonne 2009)Google Scholar argues for an increase in uncultivated land starting already in the third century in the west, which might have accelerated due to further declines in the sixth century.

83 For the originals: McCormick, ‘Rats, communications, and plague’; McCormick, ‘Tracking mass death (I)’; Sarris, ‘The Justinianic Plague.’

84 Harper, The Fate of Rome.

85 Horden, ‘Mediterranean plague’.

86 Durliat, ‘La peste du VIe siècle’.

87 Whittow, M., The Making of Byzantium, 600–1025 (Berkeley 1996), 6668CrossRefGoogle Scholar; Wickham, C., Medieval Europe (New Haven 2016), 43–4Google Scholar; Sarris, ‘The Justinianic Plague.’

88 Foss, C., ‘Syria in transition, A. D. 550–750: An archaeological approach’, Dumbarton Oaks Papers 51 (1997) 260CrossRefGoogle Scholar.

89 Bachrach, ‘Plague, population, and economy.’ This is not an uncommon position: Devroey, J-P., Économie rurale et société dans l'Europe franque: VIe - IXe siècles. T. 1: Fondements matériels, échanges et lien social (Paris 2003)Google Scholar; Devroey, ‘Catastrophe, Crise et Changement Social’, 146, 148–9.

90 Mordechai, L. and Eisenberg, M., ‘Rejecting catastrophe: The case of the Justinianic Plague’, Past & Present 244 (2019) 350CrossRefGoogle Scholar.

91 See a brief discussion in Sallares, ‘Ecology, evolution, and epidemiology of plague’, 251.

92 Walløe, L., ‘Was the disruption of the Mycenaean world caused by repeated epidemics of Bubonic Plague?’, Opusula Atheniensia 24 (1999) 121–6Google Scholar; Cunha, C. and Cunha, B., ‘Impact of plague on human history’, Infectious Disease Clinics of North America 20, no. 2 (2006) 253CrossRefGoogle ScholarPubMed, agree that the biblical epidemic (which they date to 1320 BCE) was plague. The biblical connection dates back to the nineteenth century, see McNeill, W., Plagues and Peoples (Garden City, NY 1976), 112–13Google Scholar.

93 Rasmussen, S. et al. , ‘Early divergent strains of Yersinia Pestis in Eurasia 5,000 years ago’, Cell 163, no. 3 (2015) 571–82CrossRefGoogle ScholarPubMed. Valtueña, A. A. et al. , ‘The Stone Age plague and its persistence in Eurasia’, Current Biology 27.23 (2017) 3683–3691.e8CrossRefGoogle Scholar, followed up with more research into early plague cases (naming it the Stone Age plague).

94 Spyrou, M. et al. , ‘Analysis of 3800-year-old Yersinia Pestis genomes suggests a Bronze Age origin for Bubonic Plague’, Nature Communications 9.1 (2018) 2234CrossRefGoogle ScholarPubMed.

95 Meier, M., Das andere Zeitalter Justinians: Kontingenzerfahrung und Kontingenzbewältigung im 6. Jahrhundert n. Chr (Göttingen 2003)Google Scholar; then summarized in Meier, ‘The “Justinianic Plague”’; also see the case studies on various sources in Meier, M., ‘Von Prokop zu Gregor von Tours: Kultur- und mentalitätengeschichtlich relevante Folgen der “Pest” im 6. Jahrhundert’, in Steger, F. and Jankrift, K. P., Gesundheit - Krankheit: Kulturtransfer medizinischen Wissens von der Spätantike bis in die Frühe Neuzeit (Köln 2004), 1940Google Scholar; and Meier, M., ‘Natural disasters in the chronographia of John Malalas: Reflections on their function - an initial sketch’, Medieval History Journal 10.1–2 (2007) 237–66CrossRefGoogle Scholar; or Meier, M., ‘Prokop, Agathias, Die Pest und Das, Ende‘ der Antiken Historiographie’, Historische Zeitschrift 278, no. 1 (2014) 281310Google Scholar.

96 Ubl, K., Inzestverbot und Gesetzgebung: die Konstruktion eines Verbrechens (300–1100) (Berlin 2008) 163Google Scholar.

97 See, for example, Brown, P., The Rise of Western Christendom: Triumph and Diversity, A.D. 200–1000, 10th anniversary rev. edn (Chichester 2013)Google Scholar.

98 On Mary, see for example, Shoemaker, S., Mary in Early Christian Faith and Devotion (New Haven 2016)CrossRefGoogle Scholar.

103 Note that the dataset names can be slightly different to those in the digital application.

105 See also the series of lectures on plague in Harvard's Initiative for the Science of the Human Past (webpage:

106 Stathakopoulos, Famine and Pestilence; Stathakopoulos, ‘The Justinianic Plague revisited’, remains cautious although he would move towards the maximalist interpretation over the next few years; for example, Stathakopoulos, ‘Invisible protagonists’.