Rats, communications, and plague: Toward an ecological history

Journal of Interdisciplinary History

Volumn 34, Issue 1, 2003, Pages 1-25

  McCormick, Michael ^a  

^a HARVARD UNIVERSITY   (United States)

Author keywords

[No Author keywords available]

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EID: 1842652408     PISSN: 00221953     EISSN: None     Source Type: Journal    
DOI: 10.1162/002219503322645439     Document Type: Review

References (218)

1. David E. Davis, "The Scarcity of Rats and the Black Death: An Ecological History," Journal of Interdisciplinary History, XVI (1986), 455-470.
2. David T. Dennis et al., Plague Manual (Geneva, 1999), available from http://www.who. int/emc-documents/plague/whocdscsredc992c.html, 11-12, 63.
3. e siècles)," Schweizerische Zeitschrift für Geschichte, XXIII (1973), 650-682,
4. championed the demographic impact of the pneumonic plague. See Ole J. Benedictow, Plague in the Late Medieval Nordic Countries (Oslo, 1992), 214-227,
5. for a critique of the Scandinavian historians who followed him. Its 100% mortality is indeed terrifying. Clinical data for the modern strains of Y. pestis confirm the contagiousness and lethality of the primary pneumonic form of the disease, but also qualifies the impression of it as a uniquely devastating form. See Benedictow, Plague, 25-37;
6. Dennis et al., Plague Manual, 45-46, with further references. The 1997 outbreak confirms the clinical data.
7. See Mahery Ratsitorahina et al., "Epidemiological and Diagnostic Aspects of the Outbreak of Pneumonic Plague in Madagascar," Lancet, 355 (2000), 111-113,
8. who estimate the infection rate in the contact population at 8.4%. Jean-Claude Beaucournu, "Diversité des puces vectrices en fonction des foyers pesteux," Bulletin de la Société de pathologie exotique, XCII (1999), 420.
9. Today the grass rat inhabits central Africa and the Nile river; it shares many insect parasites with the black rat, including the oriental rat flea, Xenopsylla Cheopis, the most efficient vector of Y. pestis from rodents to man. Grass rats collected in the Fayum ranked ahead of the black rat in flea counts: Mohamed L. M. Khalid, Tosson A. Morsy et al., "Studies on Flea Fauna in El Fayoum Governorate, Egypt," Journal of the Egyptian Society of Parasitology, XXII (1992), 783-799, 791 (Table 4), 796.
10. African shrew mice (Suncus murinus) also harbor the plague. Although they do not seem to transmit the disease to humans, their enzootic infection could explain how plague reservoirs sometimes outlive the temporary extinction of rat colonies. Outbreaks in Madagascar from 1991 to 1996 suggest that the plague survives in another rodent species that transmits it to new rat colonies when they spring up again: S. Laventure et al., "Épidémies de peste urbaine à Majunga, côte ouest de Madagascar," Bulletin de la Société de pathologie exotique, XCI (1998), 85-86.
11. In the United States, squirrels and prairie dogs have become reservoirs of plague; they transmit it to domestic cats, which infect their handlers. See, for example, Centers for Disease Control, "Pneumonic plague - Arizona, 1992," Journal of the American Medical Association, CCLXVIII (1992), 2146-2147.
12. See also M. J. Keeling and C. A. Gilligan, "Bubonic Plague: A Metapopulation model of a Zoonosis," Proceedings of the Royal Society of London. Biological Sciences, 267 (2000), 2219-2230.
13. The gray rat (rattus norvegicus) seems to have entered early modern Europe from the east, through the Baltic. Four or five apparently earlier remains have been assigned to this species but are still controversial - for one, on the basis of its size, the fragment of a rat femur from a fourth-century Roman well, found together with unambiguous remains of a black rat. See Gabrielle Sorge, "Ratten aus dem spätantiken Kastell Krefeld-Gellep," Provinzialrömische Forschungen. Festschrift für Günter Ulbert zum 65. Geburtstag (Espelkamp, 1995), 387-395 (with further possible cases), who hypothesizes that this gray rat, perhaps already dead, arrived with materials accompanying troops from the Roman east. Today the two kinds of rat do not normally share the same habitat. Moreover, the size of modern black rats might be a misleading criterion, since substantially larger ones are documented in medieval Corsica.
14. See Jean-Dominique Vigne et al., "Évolution ostéométrique de Rattus rattus et de Mus musculus domesticus sur de petites îles," Mammalia, LVII (1993), 85-98.
15. Serious work on the Justinianic pandemic began with Jean-Noël Biraben and Jacques Le Goff, "The Plague in the Early Middle Ages," in Robert Forster and Orest Ranum (eds.) (trans. Elborg Forster and Patricia M. Ranum), Biology of Man in History (Baltimore, 1975), 48-80;
16. Lawrence I. Conrad, "The Plague in the Early Medieval Near East," unpub. Ph.D. diss. (Princeton University, 1981).
17. Two new volumes are currently in press, Dionysios Stathakopoulos, Famine and Pestilence in the Late Roman and Early Byzantine Empire,
18. and the acts of the international conference on the Justinianic plague, Rome, 2001, edited by Lester K. Little.
19. Among studies of the medieval pandemic, three deserve special mention: Biraben, Les hommes et la peste (Paris, 1975);
20. Benedictow, Plague; La Peste Nera (Spoleto, 1994).
21. On amplifying vs. reservoir hosts, see, for example, J. D. Poland and A. M. Barnes, "Plague," in James H. Steele (ed.), CRC Handbook Series in Zoonoses: Section A. Bacterial, Rickettsial, and Mycotic Diseases (Boca Raton, 1979), I, 534-537.
22. For human blood's lower bacillus concentrations that militate against human-to-human infection through blood transferred by insect vectors, see Benedictow, Plague, 242-264;
23. Thomas Buder et al., "Yersinia Pestis Infection in Vietnam. II," Journal of Infectious Diseases, CXXXIII (1976), 493-499.
24. On fleas, see, for example, Norman Gratz, in Dennis et al., Plague Manual, 65-66;
25. Poland and Barnes, "Plague," 532, 538.
26. See Keeling and Gilligan, "Bubonic Plague," 2219-2230, for a new mathematical model, which introduces some nuance into the idea that new infections originated outside Western and Central Europe. In modern Egypt, an optimistic sounding 1% of all stored grain is reported lost.
27. For Egypt and Turkey, see M. Lund, "Commensal Rodents," in Alan P. Buckle and Robert H. Smith (eds.), Rodent Pests and Their Control (Wallingford, U.K., 1994), 23-43, 25 (Table 2.2);
28. A. N. Meyer, "Rodent Control in Practice: Food Stores," Rodent Pests and Their Control ibid., 276.
29. have not found a similar eating statistic for the slightly smaller black rat, which is a fussier eater and reportedly disappears from areas lacking its preferred foods of cereals and fruits: Lund, "Commensal Rodents," 34-35.
30. On saints' power, see the tenth- or eleventh-century Vita S. Urbani episcopi Lingonensis, XIV. Ada sanctorum Jan. III (Paris, 1863). 106, which calls the rodents both mures and sorices (cf. Fr. souris); for the date,
31. see HAGIOGRAPHIES: Sociologie et histoire de la littérature hagiographique en Occident des origines à 1550, Guy Philippart's database of Latin hagjography (http://www.fundp.ac.be/philo_lettres/histoire/h221.htm).
32. The Bollandists assign Paris, B.N., lat. 9376, the earliest ms., to the eleventh century: Bibliotheca hagiographica latina manuscripta: Index analytique des Catalogues de manuscrits hagiographiques latins publiés par les Bollandistes (http://bhlms.fltr.ucl.ac.be).
33. Graham Twigg, The Black Death: A Biological Reappraisal (London, 1984), 75-112 (insufficient rats);
34. Davis, "Scarcity of Rats" ("scarce or absent").
35. Susan Scott and Christopher J. Duncan, Biology of Plagues. Evidence from Historical Populations (Cambridge, 2001), 56-58, 108-109,
36. accept much of the reasoning of Davis and Twigg. David Herlihy, The Black Death and the Transformation of the West (Cambridge, Mass., 1997), 26-27;
37. Scott and Duncan, Biology of Plagues, 359-360;
38. Twigg, Black Death, 111-112 (including a reference to the Justmianic plague).
39. Theodoretus (ed. L. Parmentier and F. Scheidweiler), Historia ecclesiastica, 5. 39, 9-10,
40. Die griechischen christlichen Schriftsteller (Berlin, 1954; 2d ed.), XLIV, 344.4-10.
41. Oxford English Dictionary Online, s.v., suspects that rat entered European vocabulary through the Germanic languages, citing an Anglo-Saxon gloss of c. 1000; cf. Antonette di Paolo Healey (ed.), Dictionary of Old English: Old English Corpus [online] (Ann Arbor, 1998),
42. s.v. Raturus ræt, in a list of animal names, between beaver and otter. These glosses' script dates from the first half of the eleventh century: Neil R. Ker, Catalogue of Manusaipts Containing Anglo-Saxon (Oxford, 1990; 2d ed.), 1-3.
43. An eleventh-century hand glosses glis, gliris (dormouse) as ratta in a ninth-century ms.: Hartwig Mayer, Althochdeutsche Glossen: Nachträge (Toronto, 1974), 116.
44. Further research into the glosses reported by Taylor Starck and J. C. Wells, Althochdeutsches Glossenwörterbuch (Heidelberg, 1990), 2.474, might turn up an earlier occurrence. For French,
45. see Alain Rey (ed.), Dictionnaire historique de la langue française (Paris, 1998; 2d ed.), 3094.
46. First attested c. 1175, the word often means mouse in dialects. Ernst Gamillscheg, Etymologisches Wörterbuch der französischen Sprache (Heidelberg, 1969; 2d ed.), 750,
47. notes the Celtic languages. W. J. Van Douwen and J. P. N. Land (trans.), "Joannis episcopi Ephesi Syri Monophysitae Commentarii de beatis orientalibus et Historiae ecclesiasticae fragmenta," Verhandelingen der koninklijke akademie der wetenschappen, Afdeeling Letterkunde, XVIII (1889), Frag. G, 234.20-25.
48. Didier Raoult et al., "Molecular Identification by 'Suicide PCR' of Yersinia Pestis as the Agent of Medieval Black Death," Proceedings of the National Academy of Sciences of the U.S.A., XCVII (2000), 12880-12883;
49. Michel Drancourt et al., "Detection of 400-Year-Old Yersinia Pestis DNA in Human Dental Pulp," Proceedings of the National Academy of Sciences of the U.S.A., ibid., XCV (1998), 12637-12640.
50. On the early finds, see, for example, Kurt Becker, "Rattus Rattus," in Jürgen Niethammer and Franz Krapp (eds.), Handbuch der Säugetiere Europas (Wiesbaden, 1978), I, 382-400.
51. For further bibliographic material, see Frédérique Audoin-Rouzeau and Vigne, "La colonisation de l'Europe par le rat noir," Revue de paléobiologie, XIII (1994), 124-145.
52. Günther E. Thüry, "Zur Infektkette der Pest in hellenistisch-römischer Zeit," 75. Jahre Anthropologische Staatsammlung München, 1902-1977. Festschrift (Munich, 1977),
53. which I obtained only as this article went to press, also presents valuable Talmudic evidence, some of it about rattus norvegicus in Mesopotamia. Terry O'Connor, The Archaeology of Animal Bones (Stroud, 2000), 48, 50
54. (Figure 5.7) for the gnawing patterns of rattus rattus, and 2-3, 123-125 for owls' regurgitation of undigested bones and fur in the form of pellets. Apparently sterile sieve residue from Corsica produced remains of "small mammals on the order of five to ten individuals per ten liters of sediment" (Audoin-Rouzeau and Vigne, "La colonisation," 145, n. 3;
55. O'Connor, Archaeology, 28-35).
56. Sorge, "Ratten," adds a few more sites.
57. Since 1994, more than thirty new finds from about twenty ancient and medieval sites have come to light. For the first-century A.D. Roman rats near Amsterdam and in Burgundy, see Neli Gordijn-Vons, "Een voorbericht over dierbotvondsten van Romeins-Velsen uit de eerste helft van de eerste eeuw," Westerheem, XXVI (1977), 125 (Table 5);
58. Sébastien Lepetz et al., "Nouvelles observations du rat noir (Rattus rattus) dans la moitié nord de la France à la période gallo- romaine," Revue archéologique de Picardie (1993), 173-174.
59. For second-century and medieval rats from England, see Alison Locker, "The Mammal, Bird and Fish Bones," in David S. Neal et al. (eds.), Excavation of the Iron Age, Roman, and Medieval Settlement at Gorhambury, St. Albans (London, 1990) 211 (Table 23).
60. For late Roman rats in southern Belgium, see Jan De Coninck et al., "L'abri de la Sigillée. VI," Anthropologie et Préhistoire, CI (1990), 39-45;
61. in a Roman Rhine fort, Sorge, "Ratten"; in southern Italy,
62. Salvatore Scali, "Observations on the Faunal Remains from the Territory of Metaponto," The Territory of Metaponto 1981-1982 (Austin, 1983) (unpaginated);
63. in central Italy, Michael M[a]cKinnon, "Animal Bone Remains," in David Soren and Noelle Soren (eds.), A Roman Villa and a Late Roman Infant Cemetary: Excavation at Poggio Gramignano Lugnano in Teverina (Rome, 1999), 539 (Table 1), 537, 555-556.
64. For seventh- and eighth-century Syria, see Achille Gautier, "La faune de quelques maisons d'Apamée," in Janine Balty (ed.), Apamée de Syrie. Bilan des recherches archéologiques, 1973-1979 (Brussels, 1984), 305-360.
65. For late Roman and medieval rats at a papal farm north of Rome, see Anthony King, "Mammal, Reptile and Animal Bones," in T. W. Potter and A. C. King (eds.), Excavations at the Mola di Monte Gelato (London, 1997), 383-403.
66. For many medieval sites, see Antonio Tagliacozzo, "I reperti faunistici," in Carlo Pavolini (ed.), Caput Africae (Rome, 1993), 258;
67. Philip L. Armitage, "Unwelcome Companions: Ancient Rats Reviewed," Antiquity, LVIII (1994), 234,
68. which discusses the "Byzantine" rat remains reported by M. Stanzel from the excavations of the Artemis-Apollo temple at Kalopedi, in Boeotia; Hans Reichstein, "Über Kleinsäuger aus Burg Bodenteich," Bonner zoologischer Beiträge, XLVI (1990), 359-366;
69. Hans Reichstein, idem, "Die wildlebenden Säugetiere von Haithabu," Berichte über die Ausgrabungen in Haithabu, XXX (1991), 31-33,
70. at least ten "early medieval" rats, with no stratigraphy; Bengt Wigh, Animal Husbandry in the Viking Age Town of Birka and Its Hinterland (Stockholm, 2001), 54 (Table 10), 125-126,
71. from five different strata dating between c. 810-830 to c. 950. For thirteenth-century Muslim Portugal, see A. Morales and J. Rodriguez, "Black Rats (Rattus rattus) from Medieval Mertola," Journal of Zoology (London), 241 (1997), 623-642,
72. with valuable methodological reflections. For twelfth-century Namur, see W. Van Neer and A. Lentacker, "Restes fauniques provenant de trois fosses d'aisances du Grognon à Namur," in J. Plumier and M. H. Corbiau (eds.), Quatrième journée d'archéologie namuroise (Namur, 1996), 89-104, Tableau 3;
73. 1300 and the fourteenth-fifteenth century, B. De Cupere and I. Boone, "Le matériel faunique du château des comtes à Namur. Résultats préliminaires," in J. and S. Plumier-Torfs and C. Duhaut (eds.), Huitième journée d'archéologie namuroise (Rochefort, 2000), 11-16.
74. For late medieval Brabant, see Anton Ervynck et al., "Dierlijke resten," in idem, De "burcht" te Londerzeel (Zellik, 1994), 155, cf. 181, 186, 211 (Tabel J), 213 (Tabel L). For the other ancient and medieval rat finds, see nn. 10 and 12 below. As this article went to press, John Clark kindly put me in touch with Kevin Rielly (both from the Museum of London), who generously alerted me to three unpublished eleventh-century rat finds, from the City of London (Bull Wharf, No. 1 Poultry and Guildhall sites), as well as another one of Saxon date from a pit excavated at the National Gallery Basement site (NGA87). These four finds are not included in my totals.
75. Audoin-Rouzeau and Vigne, "La colonisation," 126.
76. For Malaysia, see Ronald M. Nowak (ed.), Walker's Mammals of the World Online (Baltimore, 1997; 5th ed.), s.v. rattus.
77. For karyotypes and diffusion, see Jürgen Niethammer, "Zur Taxonomie und Ausbreitungsgeschichte der Hausratte (Rattus rattus)," Zoologischer Anzeiger, 194 (1975), 405-415,
78. confirmed by R. Libois et al., "Les populations de rats noirs insulaires de l'ouest de l'Europe," Vie et milieu, XLVI (1996), 213-218.
79. On ancient trade with the region and the ports called Muziris and Nelkynda, see Lionel Casson, The Periplus Maris Erythraei (Princeton, 1989), 21-25.
80. See, for example, E. Bresciani, "Persia," Oxford Encyclopedia of Ancient Egypt (Oxford, 2001), III, 37;
81. J. S. Holladay, Jr., "Pithom," ," Oxford Encyclopedia of Ancient Egypt ibid., 50-53;
82. J. S. Holladay, idem, "Wadi Tumilat," in Kathryn A. Bard and Steven B. Shubert (eds.), Encyclopedia of the Archaeology of Ancient Egypt (London, 1999), 878-881;
83. P. J. Sijpesteijn, "Der Potamos Traianos," Aegyptus, XLIII (1963), 70-83.
84. The alternative translation of "Myos Hormos" as "Port of the Mussel" is widely accepted, but "mussel" is a secondary meaning of the word mys. For the site, see D. Whitcomb, "Quseir el-Qadim," Encyclopedia of the Archaeology of Ancient Egypt, 658-660;
85. for the Roman rats, Angela von den Driesch and Joachim Boessneck, "A Roman Cat Skeleton from Quseir on the Red Sea Coast," Journal of Archaeological Science, X (1983), 210-211.
86. From the rich cave deposits on Monte di Tuda, c. 10 km from the sea, see Vigne and H. Valladas, "Small Mammal Fossil Assemblages as Indicators of Environmental Change in Northern Corsica during the last 2500 Years," Journal of Archaeological Science, XXIII (1996), 199-215.
87. For the early but isolated claims, see Audoin-Rouzeau and Vigne, "La colonisation," 129 (Tableau 1, nos. 0-5), 132.
88. For the single fragment tentatively assigned to the seventh-sixth century B.C. from an Adriatic site, see Tassos Kotsakis and Elena Ruschioni, "I microvertebrati di un insediamento dell'Età del Ferro presso Tortoreto," Rendiconti della Accademia nazionale dei Lincei. Classe di scienze fisiche, matematiche e naturali, LXXVI (1984), 295-304.
89. Audoin-Rouzeau and Vigne, "La colonisation," nos. 20, 24.
90. L. Lortet and C. Gaillard, "La faune momifiée de l'ancienne Égypte," Archives du Muséum d'histoire naturelle de Lyon, VIII (1903), 39-40, discovered a number of partially digested black rats in mummified birds of prey supplied by the Egyptologist Gaston Maspero (1846-1916). Lortet does not explicitly date them but states that they came from Giza and Kom Ombo in Upper Egypt. That he dates his bird mummies from Giza as Ptolemaic and, "maybe, XXVI dynasty," that is, 663-525 B.C. (113-114), suggests that some of the Giza mummies might have been earlier, but that most, if not all, were Ptolemaic. Those from Kom Ombo were Roman.
91. Libois et al., "Les populations";
92. G. Cheylan et al., "Distribution of Genetic Diversity within and between Western Mediterranean Island Populations of the Black Rat Rattus rattus," Biological Journal of the Linnean Society, LXIII (1998), 393-408.
93. For amphoras at Tell el-Maskhuta, see Holladay, "Pithom," 53.
94. Excluded from my sixty-five contexts are the seven northern finds for which Audoin-Rouzeau raises the possibility of an intrusive deposit: nos. 12 and 32, Ersigen-Murain and Vendeuil-Caply (both first century); 16, Lorch-Lauriacum (fifth century); 19, Magdalensberg, (first century B.C. - first century A.D.), 21; Muensterberg Breisach and 29, Sponeck (both fourth century); and 58, the poorly dated Unterregenbach (seventh-eleventh century). Ten finds are dated to a range that extends beyond the eighth century, including one that might also come from the ninth century; six might be as late as the tenth century (including the new data from Haithabu) (see n. 7 above); and one context each is dated eighth to thirteenth and seventh to sixteenth century. Audoin-Rouzeau and Vigne, "La colonisation," 129 (Tableau 1): nos. 11, Carthage (first to fifth century A.D.); 35, Zembra (seventh century); 26, Quseir el-Qadim (first or second century, six rats); the Apamaea finds (sixth to seventh and seventh to eighth century); the mummified birds in nn. 7 and 10.
95. On rats' limited range, see, for example, Biraben, Les hommes et la peste, I, 17,
96. relying essentially on J. F. D. Shrewsbury, A History of the Bubonic Plague in the British Isles (Cambridge, 1970), 7-16,
97. which seems to draw on R. A. Donaldson, The Rat (Philadelphia, 1924);
98. M. A. C. Hinton, Rats and Mice as Enemies of Mankind (London, 1918).
99. have not found much empirical study of black rat movements in the wild. Radio tracking of three rats in a temperate wooded area near Sydney, Australia, indicated a home range of 0.76 to 0.3 ha (100 x 100 m sq) over a two- or three-day period. See Michelle P. G. Cox et al., "Use of Habitat by the Black Rat (Rattus rattus) at North Head, New South Wales," Austral Ecology, XXV (2000), 380;
100. cf., for example, J. G. Innes and J. P. Skipworth, "Home Ranges of Ship Rats in a Small New Zealand Forest," New Zealand Joumal of Zoology, X (1983), 99-110.
101. Trapping and release experiments point to restricted home ranges. Over two months, G. F. Petrie et al., "A Report on Plague Investigations in Egypt," Reports and Notes of the Public Health Laboratories, Cairo, V (1923), 1-114,
102. noted only one rat moving beyond a restricted (but ill-defined) home range in a block of houses in El Motiya, Egypt. That rat traveled 250 m. As the authors note, because this experiment coincided with a food shortage, breeding, and plague, movements may well have been maximal. However 159 of the 305 marked rats were not captured again, so we know nothing about their movements (77). The thorough observations of Hans Joachim Telle, "Beitrag zur Kenntnis der Verhaltensweise von Ratten," Zeitschrift für angewandte Zoologie, LIII (1966), 144-145,
103. indicate that 181 of 212 German black rat packs remained within a territory that included their food source; the only territory for which a size is given was a 12 x 4 m roof (150, Abb. 4). Warren F. Pippin's year-long study on a tropical island, "The Distribution and Movement of Roof Rats on Mona Island, West Indies," Journal of Mammalogy, XLII (1961), 344-348,
104. does not fundamentally contradict the picture; rats released where they were captured were never recaptured more than 50 yds (45.5 m) away. Most rats released at various distances from the point of capture made their way back, suggesting a strong sense of territoriality. Some of those "homing" rats, however, traveled well over 1,000 yards (914 m) in a day or two; one covered 8,700 yds (7.95 km) in 48 hours. Four rats seem to have chosen to travel elsewhere, at distances of 950, 900, 600 (one to two days) and 350 yds (thirteen days). Finally, Telle, "Beitrag," 145, observed that unlike northern rats, Mediterranean rats made burrows, did not shun wet environments, and nested as far as 300 to 400 m from food sources; an ecological gradient may also be in play at Mona Island. According to my assumptions, estimated life span of rats in the wild is 2 to 2.5 years. Rats begin bearing after c. 100 days of life, and are pregnant for about twenty-two days. They may average four or five pregnancies during their lifetime. See Becker, "Rattus rattus," 394-395.
105. Conditions may vary according to the ecological circumstances. E. W. Bentley, "The Distribution and Status of Rattus rattus L. in the United Kingdom in 1951 and 1956," Journal of Animal Ecology, XXVIII (1959), 302-303;
106. Colin Matheson, "A Survey of the Status of Rattus Rattus and Its Subspecies in the Seaports of Great Britain and Ireland," ," Journal of Animal Ecology, ibid., VIII (1939), 86-89.
107. Telle, "Beitrag," 144-145.
108. Armitage, "Unwelcome Companions," 234,
109. mentions a rat discovered in the Roman bath at Mirebeau. Lepetz et al., "Nouvelles observations," 173-174.
110. J. C. Lowe and S. Moryadas, The Geography of Movement (Boston, 1975), 150-157.
111. For comments on pottery circulation between Monte Gelato and Rome, see Helen Patterson, "The Early Medieval and Medieval Pottery," in Potter and King (eds.), Excavations, 374,
112. and, in general, Peregrine Horden and Nicholas Purcell, The Corrupting Sea (Oxford, 2000), 110-112.
113. Audoin-Rouzeau and Vigne, "La colonisation," 135;
114. Sharon J. Downes et al., "Variation in the Use of Corridors by Introduced and Native Rodents in South-Eastern Australia," Biological Conservation, LXXXII (1997), 379-383.
115. See, in general, McCormick, Origins of the European Economy: Communications and Commerce, A.D. 300-900 (New York, 2001), 67-77.
116. For a stimulating, positive appraisal of the economics of road transport, see Ray Laurence, "Land Transport in Roman Italy," in Helen Parkins and Christopher Smith (eds.), Trade, Traders and the Ancient City (London, 1998), 129-148.
117. McCormick, "Bateaux de vie, bateaux de mort," Morfologie sociali e culturali in Europa fra tarda antichità e alto medioevo (Settimane di studi del Centro italiano di studi sull'alto medioevo, 45) (Spoleto, 1998), 58-61.
118. For references to Rhone and Alps transports, see McCormick, Origins, 77-80.
119. Walter Janssen, "Reiten und Fahren in der Merowingerzeit," in H. Jankuhn et al. (eds.), Der Verkehr (Abhandlungen der Akademie der Wissenschaften in Göttingen, Philologisch-historische Klasse, 3rd series, 180) (Göttingen, 1989), 198-202.
120. For Augustus' road to the Little St. Bernard, see Raymond Chevallier, Les voies romaines (Paris, 1997), 213;
121. M. Hudry, "Trace et trafic d'une voie romaine transalpine: section Petit-Saint-Bernard-Albertville," Actes du colloque international sur les cols des Alpes (Bourg en Bresse, 1969), 109.
122. Most authors believe that the Great St. Bernard, however, was suited only for mule trains. See R. Fellmann, "Le strade romane attraverso il Massicio Alpino della Svizzera," in M. Pavan and G. Rosada (eds.), La 'Venetia' nell'area padano-danubiana (Padua, 1990), 370-371.
123. Dissenting is P. Hunt, "Summus Peninus on the Grand St Bernard Pass," Journal of Roman Archaeology, XI (1998), 265-274.
124. Richard Bulliet, The Camel and the Wheel (Cambridge, Mass., 1975);
125. Roger S. Bagnall, "The Camel, the Wagon and the Donkey in Later Roman Egypt," Bulletin of the American Society of Papyrologists, XXII (1985), 1-6.
126. In Senegal, the distribution of rat colonies has changed since trains supplanted riverboats as the main line of communications with Mali. See J. M. Duplantier et al., "Répartition actuelle du rat noir (Rattus rattus) au Sénégal: Facteurs historiques et écologiques," in Michel Le Berre and Louis Le Guelte (eds.), Le rongeur et l'espace (Saint-Etienne, 1991), 345.
127. Becker, "Rattus rattus," 393-394;
128. McCormick, "Bateaux de vie, bateaux de mort";
129. Michael F. Hendy, Studies in the Byzantine Monetary Economy (Cambridge, 1985), 294-296, 603-608.
130. See Poland and Barnes, "Plague," 532.
131. Keeling and Gilligan, "Metapopulation dynamics of bubonic plague," Nature, CDVII (2000), 905.
132. The point about medieval Europe's rat colonies is based on Audoin-Rouzeau and Vigne, "La colonisation," 128-131 (Table 1), and the relevant material in n. 7.
133. Thus, Morales and Rodriguez, "Black Rats (Rattus rattus) from Medieval Mertola," 624-632,
134. cautiously formulate the age distribution for a group of thirteenth-century rats, arguing that it shows an attritional (rather than catastrophic) age structure. See Keeling and Gilligan, "Bubonic Plague,"
135. and Keeling and Gilligan, idem, "Metapopulation Dynamics."
136. Although now dated, the classic statement of the cemetery method is Peter Donat and Herbert Ullrich, "Einwohnerzahl und Siedlungsgrösse der Merowingerzeit," Zeitschrift für Archäologie, V (1971), 234-265.
137. For a recent application, see, for example, Klaus Georg Kokkotidis, "Belegungsablauf und Bevölkerungsstruktur auf dem alamannischen Gräberfeld von Fridingen an der Donau in Südwestdeutschland," Fundberichte aus Baden-Württemberg. Aufsätze, XX (1995), 737-801.
138. For a far-reaching critique of similar methods for the Roman empire, see Walter Scheidel, "Roman Age Structure: Evidence and Models," Journal of Roman Studies, XCI (2001), 1-26.
139. The strata were dated c. 450-475, c. 475-525, and c. 525-699, respectively. See A. C. King, "Mammiferi," in Paul Arthur (ed.), Il complesso archeologico di Carminiello ai Mannesi, Napoli (Galatina, 1994), 376 (Tabella 39), 387-388, 405.
140. Arthur, Il complesso archeologico ibid., discusses the dating, although the differing precision needs to be taken into account, in the introduction ("Sintesi delle principali attività per fasi," 73-75) and the conclusion ("Conclusioni," 433-435).
141. Becker, "Rattus rattus," 387-390;
142. Grzimek's Encyclopedia of Mammals (New York, 1990), 3.167.
143. For the broad late Roman demographic trends, see, for example, McCormick, Origins, 30-41.
144. Becker, "Rattus rattus," 394-396;
145. McCormick, "Bateaux de vie, bateaux de mort," 37-43,
146. with further references; Jean-Pierre Devroey, "La céréaliculture dans le monde franc," L'ambiente vegetale nell'alto medioevo (Settimane di studio del Centro italiano di studi sull'alto medioevo, 37) (Spoleto, 1990), 231-232.
147. On sanitation in Roman cities, see A. Scobie, "Slums, Sanitation and Mortality in the Roman World," Klio, LXVIII (1986), 399-433.
148. On Roman waste and the implications for rats, see Thüry, Die Wurzeln unserer Umweltkrise und die griechisch-römische Antike (Salzburg, 1995), 23-26, 77, n. 61;
149. Thüry, idem, Müll und Marmorsäulen (Mainz, 2001).
150. For Roman trash law, see, in particular, Digesta, 43, 10,
151. in Theodor Mommsen and Paul Krueger (eds.), Corpus iuris avilis (Berlin, 1922), I, 733;
152. cf. Digesta, 9, 3, 1-7,
153. Digesta, ibid., 163-164;
154. Digesta, 44, 7, 5,
155. Digesta, ibid., 765, on injuries caused by refuse thrown out of windows onto passersby.
156. Arthur, "Sintesi," 73.
157. Bentley, "Distribution," 303-304.
158. Yersin quoted in Thomas Butler, Plague and Other Yersinia Infections (New York, 1983), 17.
159. Petrie et al., "Report," 16.
160. For broad streets, rats, and Japanese cities, see the abstract of Tatsuo Yabe, "Changes in Species Composition of the Norway Rat (Rattus norvegicus) and the Roof Rat (R. rattus) in Urban Area [in Japanese]," Medical Entomology and Zoology, XLVIII (1997), 285-294,
161. according to BIOSIS Previews online. James Russell, "Transformations in Early Byzantine Urban Life," 17th International Byzantine Congress: Major Papers (New Rochelle, 1986), 144;
162. Jean Durliat, "L'approvisionnement de Constantinople," in Cyril Mango and Gilbert Dagron (eds.), Constantinople and Its Hinterland (Aldershot, 1995), 30-32.
163. For Alexandria, see Naguiba F. Loutfy et al., "Study on Rodents Infestation in Alexandria and Prevalence of Trichinella Spiralis Infection among Them," Journal of the Egyptian Society of Parasitology, XXIX (1999), 897-909, according to BIOSIS Previews online.
164. For London (without explicit mention of rats), see Twigg, Black Death, 110-111.
165. For recent plague epidemics near a city market and its garbage, see Pascal Boisier et al., "Epidemiologic Features of Four Successive Annual Outbreaks of Bubonic Plague in Mahajanga, Madagascar," Emerging Infectious Diseases, VIII (2002), 314.
166. MacKinnon, "High on the Hog: Linking Zooarchaeological, Literary, and Artistic Data for Pig Breeds in Roman Italy," American Journal of Archaeology, CV (2001), 649-673.
167. According to Erich Woehlkens, Pest und Ruhr im 16. und 17. Jahrhundert (Hanover, 1954), 72-74, only one-quarter of smith or cooper enterprises were hit by plague in 1597, whereas 84% of other shops were affected.
168. Although rodent hearing is highly sensitive, ultrasound devices seem fairly ineffective against them, in part because their initial aversion is overcome by habit. If the German case is valid, the factors of rat deterrence there may have been more complex, involving the availability of alternative habitats in the vicinity. On rodent hearing, see D. W. Macdonald and M. G. P. Fenn, "The Natural History of Rodents: Preadaptations to Pestilence," in Buckle and Smith (eds.), Rodent Pests, 6.
169. On their habituation to noise, see Smith, "Rodent Control Methods: Non-chemical and Non-lethal Chemical," in Rodent Pests, ibid., 113-114 (with Table 5.1).
170. Becker, "Rattus rattus," 388.
171. Vijay K. Saxena and T. Verghese, "Ecology of Flea-Transmitted Zoonotic Infection in Village Mamla, District Beed," Current Science (Bangalore), LXXI (1996), 800-802, from the abstract in BIOSIS Previews online.
172. See also S. Shivaji et al., "Identification of Yersinia Pestis as the Causative Organism of Plague in India as Determined by 16S rDNA Sequencing and RAPD-based Genomic Fingerprinting," FEMS Microbiology Letters, 189 (2000), 247-252,
173. confirming the Y. Pestis diagnosis. Glanville Downey, "Earthquakes at Constantinople and Vicinity, A.D. 342-1454," Speculum, XXX (1955), 598-599.
174. For the 558 plague, see Stathakopoulos, "Crime and Punishment: The Plague in the Byzantine Empire," forthcoming in the volume to be edited by Little, cited in n. 3.
175. On environment and vectors, see Duane J. Gubler et al., "Climate Variability and Change in the United States: Potential Impacts on Vector and Rodent-Borne Diseases," Environmental Health Perspectives, CIX, Supplement 2 (2001), 223-233 (228-229 on plague in particular).
176. Robert R. Parmenter et al., "Incidence of Plague Associated with Increased Winter-Spring Precipitation in New Mexico," American Journal of Tropical Medicine and Hygiene, LXI (1999), 814-821.
177. Unless precipitation is equal throughout an entire region, the correlation between increased precipitation and plague holds better at the local than the regional level. Observations from Africa suggest that the precipitation-rodent link extends beyond New Mexico's arid conditions. See Rhodes H. Makundi, "Annual Changes of Reproduction in Rodents in the Western Usambara Mountains, North-East Tanzania, "Journal of African Zoology, CIX (1995), 15-21;
178. cf. Afework Bekele and Herwig Leirs, "Population Ecology of Rodents of Maize Fields and Grasslands in Central Ethiopia," Belgian Journal of Zoology, CXXVII (1997), 39-48, although there were few black rats in this last sample.
179. For Constantinople and Persia, see Johannes Koder, "Climatic Change in the Fifth and Sixth Centuries?" in Pauline Allen and Elizabeth M. Jeffreys (eds.), The Sixth Century: End or Beginning? (Brisbane, 1996), 275-276.
180. For the Po, see Procopius, Bella, 6, 28, 3-6,
181. in J. Haury and G. Wirth (eds.), Opera omnia (Leipzig, 1963), II, 275.16-276.12.
182. For Rome's fading sunlight, see Procopius, Bella, 4, 14, 5-6, 1.482.19-483.5;
183. Cassiodorus, Variae, 12, 25,
184. in Å. J. Fridh (ed.), Corpus christianorum, series latina, XCVI (1973), 492-494,
185. makes clear that the previous year had produced an exceptionally good harvest in Italy; drought accompanied the obscuring of the sun for eighteen months. See Amir Harrak (trans.), The Chronicle of Zuqnīn (Toronto, 1999), 87.
186. Koder, "Climatic Change?"
187. For the Syrian snow, see, for example, Chronicle of Zuqnīn n, 89.
188. Joel D. Gunn, "A.D. 536 and Its 300-Year Aftermath," in idem, The Years without Summer: Tracing A.D. 536 and Its Aftermath (Oxford, 2000), 12.
189. The Greenland ice cores appear to argue against a volcanic origin, according to Mike Baillie, Exodus to Arthur: Catastrophic Encounters with Comets (London, 2000; 2d ed.), 8, 65-68 (on the stress displayed by European trees in 536 and 540/1).
190. For solar emissions, see Paul Farquharson, "Byzantium, Planet Earth, and the Solar System," in Allen and Jeffreys (eds.), Sixth Century, 263-268.
191. David Keys, Catastrophe: An Investigation into the Origins of the Modern World (London, 1999), 18-19, argues that drought followed by precipitation most favors plague outbreak, citing unspecified research "monitored" by the Centers for Disease Control. Hence, for him, the Justmianic plague was caused by infected rodents migrating from their usual African habitat because of the event of 536. The chains of causality are likely more complex; it is not certain that Y. pestis originated in Africa.
192. See McCormick, "Toward a Molecular History of the Justinianic Pandemic" (forthcoming).
193. King, "Mammiferi," 387-388 (Tabella 39).
194. Although some say that rats are too big for cats, see P. D. Meek and B. Triggs, "The Food of Foxes, Dogs and Cats on two Peninsulas in Jervis Bay, New South Wales," Proceedings of the Linnaean Society of New South Wales, CXX (1998), 117-127, 124;
195. von den Driesch and Boessneck, "Roman Cat Skeleton," 210-211,
196. for the six rats discovered in a mummified Roman cat's digestive tract. For fleas and country cats, see J. H. Yvinec et al., "Premiers apports archéoentomologjques de l'étude des Puces," Bulletin de la Société entomologique de France, CV (2000), 422.
197. Vigne and Valladas, "Small Mammal Fossil."
198. Modern Italian owl diets conform to Roman and medieval owl droppings in Corsica, showing that rattus rattus is a favorite food for certain owls. See, for example, Dario Capizzi et al., "Feeding Habits of Sympatric Long-Eared Owl Asio Otus, Tawny Owl Strix Aluco and Barn Owl Tyto Alba in a Mediterranean Coastal Woodland," Acta Ornithologica [Warsaw], XXXIII (1998), 85-92.
199. New Zealand foxes (Vulpes vulpes) feed opportunistically and abundantly on black rats. See Meek and Triggs, "Food," esp. 120 (Table 1). Weasels are famous rat hunters.
200. Armitage, "Unwelcome Companions," 233-234;
201. Davis, "Scarcity of Rats," 456-457;
202. Twigg, Black Death, 99-100, 124;
203. G. Key et al., "Ship Rats Rattus rattus on the Shiant Islands, Hebrides, Scotland," Journal of Zoology [London], CCXLV (1998), 228-233.
204. The rats in the Macquarie Islands have found a niche at the base of grass plants. They have adapted to distinctly non-Mediterranean conditions; temperatures in their burrows ranged from lows of 4.8°C (cool day) and 8.7°C (warm day) to highs of 5.0°C (cool day), and 9.2°C (warm day). See T. Pye et al., "Distribution and Habitat Use of the Feral Black Rat (Rattus rattus) on Subantarctic Macquarie Island," Journal of Zoology, CCXLVII (1999), 435-436.
205. For the most certain evidence of rats in the medieval north, see Audoin-Rouzeau and Vigne, "La colonisation," 129-130 (Table 1), nos. 41-42, 44-46, 48, 51, 56, 61, from France, Belgium, Germany, England, and Sweden.
206. For the new finds in Viking-age Birka, see Wigh, Animal Husbandry, 54 (Table 10).
207. For northwestern Europe, see Audoin-Rouzeau and Vigne, "La colonisation," 130 (Table 1),
208. no. 53, where they qualify the find with a question mark. The authors also mention Boulogne, Portchester, York, Southampton, and Paris (137). At York, rats apparently were introduced c. 175-250 A.D. Unlike mice, they are absent from eighth-century deposits, reappearing under the Vikings in the late ninth century. See O'Connor, "Bones from 46-54 Fishergate," in idem, The Animal Bones (The Archaeology of York, 15.4) (Dorchester, 1991), 256-258.
209. For the "Northern Arc," see O'Connor, idem, Archaeology, 157.
210. For Birka, see Wigh, Animal Husbandry, 54 (Table 10), 125-126.
211. For York, see O'Connor, "Bones." The Dorestad absence is not significant; sieving came late there. The key mesh was large (4 mm), and only 6 g of wild small mammal remains were identified from 7801 of soil.
212. See Wietske Prummel, Excavations at Dorestad, 2. Early Medieval Dorestad: An Archaeozoological Study (Amersfoort, 1983), Table 20, 245-246;
213. 41-42 (collection methods). For the fifteen other contexts, see Audoin-Rouzeau and Vigne, "La colonisation," nos. 41-42, 45-46, 49, 52, 55, 57, 60-63, 66,
214. and Reichstein's report of the finds at Haithabu and in the castle of Bodenteich, n. 7. For the rats' return, see, for example, Wigh, Animal Husbandry, 125-126;
215. O'Connor, "Pets and Pests in Roman and Medieval Britain," Mammal Review, XXII (1992), 108.
216. Keeling and Gilligan, "Bubonic plague," 2219-2230,
217. show the model for a rat population of 60,000. Keeling and Gilligan, Idem, "Metapopulation Dynamics";
218. Elizabeth Matisoo-Smith et al., "Patterns of Prehistoric Human Mobility in Polynesia Indicated by mtDNA from the Pacific Rat," Proceedings of the National Academy of Sciences of the United States of America, XCV (1998), 15145-15150.