Zeitschrift für interdisziplinäre Geschichte, L:2 (Herbst, 2019), 171–185.

Das 50. Jahr: Sonderaufsatz 3

Anne Hardy
The Under-Appreciated Rodent: Harbingers of
Plague From the Middle Ages to the Twenty-
First Century The social and interdisciplinary history of
morbidity and mortality, of death and disease, is a relatively re-
cent phenomenon, emerging in the 1970s with the Bulletin of the
Social History of Medicine (1970)—reborn as Social History of
Medicine in 1987—roughly coeval with the Journal of Interdisci-
plinary History. Shrewsbury’s magisterial History of Bubonic Plague
in the British Isles was also published in 1970, and Lebrun’s Les
hommes et la mort en Anjou aux 17ième et 18ième sièçles, which ap-
peared in 1971, arguably set the study of death and disease on a
new track. In the years following, the historiography of the Black
Death has ﬂourished, and the history of morbidity and mortality has
become an important area of historical expertise in its own right.1
In this literature, the agents of disease themselves, rather than
their involvement with, and impact on, human societies, have been
conspicuously absent. Zinsser’s classic Rats, Lice and History (Boston,
1935), Zum Beispiel, centered on Rickettsia prowazeki, the causal
organism of typhus, not on the rat itself. Among the rich and varied
scholarly contributions to the pages of the JIH, four articles concern
the medieval Black Death, and three treat plague in other centuries.
In the ﬁrst group, the ones by Davis and McCormick downplay the
importance of rats. Davis doubted the role of the black rat, arguing

Anne Hardy is Honorary Professor of the History of Public Health, London School of
Hygiene & Tropical Medicine. She is the author of The Epidemic Streets: Infectious Disease
and the Rise of Preventive Medicine, 1856–1900 (New York, 1993); Salmonella Infections, Netzwerke
of Knowledge and Public Health in Britain, 1880–1975 (New York, 2015); “A New Chapter in
Medical History,” Zeitschrift für interdisziplinäre Geschichte, XXXIX (2009), 349–359.

© 2019 vom Massachusetts Institute of Technology und The Journal of Interdisciplinary
Geschichte, Inc., https://doi.org/10.1162/jinh_a_01408

I am grateful to Robert Rotberg for coining the phrase “the under-appreciated rodent.”
1
John F. D. Shrewsbury, History of Bubonic Plague in the British Isles (New York, 1970); François
Lebrun, Les hommes et la mort en Anjou aux 17ieme et 18ieme siècles: essai en demographie et psy-
chologie historique (Paris, 1971).

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that it is a shy, ship-dependent, and diurnal rodent that existed in
small numbers in the fourteenth century, and McCormick, working
with improved archaeological techniques after the revolution in
modern technologies, challenged the view that ship-borne rats dis-
tributed the disease through Europe, arguing instead for land-based
travel via grain wagons.2

Dean et al. recently argued, based on a study of demographic
data from nine European cities, that a human parasite model is a
much better ﬁt for the pattern of the fourteenth-century outbreak.
Jedoch, Stenseth, director of the Centre for Ecological and
Evolutionary Synthesis at the University of Oslo, who contributed
to the study of Dean et al., also maintained that gerbils, not rats, Sind
to blame for the Black Death, although gerbils, which are native to
Africa, Asien, and India, did not arrive in Europe until the nineteenth
Jahrhundert. The ﬁrst identiﬁcation of rats as the agents of plague transmis-
sion came after the 1894 Hong Kong outbreak, which prompted the
British public-health authorities to monitor plague cases on ships. In
Die Vereinigten Staaten, San Francisco’s Board of Health recommended the
rat-prooﬁng of buildings as a precaution against plague after the de-
termination that ship rats had infected the local rodent population
with Yersinia pestis c. 1900. Taking Davis’ and McCormick’s Black
Death contributions as a starting point, this article stands in oppo-
sition to the recent historiography and focuses instead on the rat as
an under-appreciated contributor to human history.3

2 The articles concerned with the Black Death in the JIH are David E. Davis, “The Scarcity of
Rats and the Black Death: An Ecological History,” XVI (1986), 455–470; Michael McCormick,
“Rats, Kommunikation, und Pest: Toward an Ecological History,” XXXIV (2003), 1–25;
John Theilmann and Frances Cate, “A Plague of Plagues: The Problem of Plague Diagnosis
in Medieval England,” XXXVII (2007), 371–393; Sharon DeWitte and Philip Slavin, “Between
Famine and Death—England on the Eve of the Black Death—Evidence from Paleopathology
and Manorial Accounts,” XLIV (2013), 37–60.
3 Katharine R. Dean et al., “Human Ectoparasites and the Spread of Plague in Europe during
the Second Pandemic,” Proceedings of the National Academy of Sciences, available at www.pnas.org/
content/early/2018/01/09/1715640115 (accessed April 1, 2019). For a counterargument, see San
Woo Park et al, “Human Ectoparasite Transmission of the Plague during the Second Pandemic Is
Only Weakly Supported by Proposed Mathematical Models,” ebenda., available at www.pnas.org/
content/115/34/7892. For a history of the view that rats did not play a causal role, see Monica
Grün, “On Learning How to Teach the Black Death,” available at https://www.academia.edu/
36171431/On_Learning_How_To_Teach_the_Black_Death_2018 (accessed April 1, 2019). Für
Nils Christian Stenseth et al., siehe https://www.history.com/news/scientists-blame-gerbils-not-
rats-for-the-black-death (abgerufen im Juli 10, 2019); Vernon B. Link and Theodore M. Bauer, A
History of Plague in the United States, Public Health Monograph 26 (Washington, D.C., 1955), 10.

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| 173

Plagues do not belong solely to the past; emerging and re-
emerging infections have become an important strand of medical
interest in the years since 1990. New infections either arose or
were identiﬁed on several occasions following the recognition of
the Marburg virus in 1967, but the general view remained that se-
rious infectious diseases were a thing the past. The shock arrival of
HIV/AIDS as a major epidemic in the 1980s marked an ominous step
in a new progression that threatened to surpass the virulence of the
largely controlled infections that predated it. Im November 1993,
Berkelsman and Hughes of the Centers for Disease Control pub-
lished an editorial in Annals of Internal Medicine that threw into
question the notion that infectious diseases were no longer a
threat. Less than a year later, the sudden explosion of epidemic bu-
bonic plague in the Indian states of Maharashtra and Gujarat in
September 1994 underscored the point, seeming to mark the re-
turn of a globally legendary disease. Since India had not conﬁrmed
any human case of plague for decades, the event acted as a catalyst
for concerns about existing vulnerabilities around the world. Seit
1994, bubonic plague and its infamous perpetrator, the rat, have
remained a constant presence in the consciousness of all those
who study and treat infectious diseases.4

The Black Death of 1347 Zu 1353 stands out in history as the
quintessential emergent infection, appearing almost out of no-
where and sweeping lethally across the European world to become
the stuff of legend. Yet although Davis and McCormick sought
effectively to dismiss the role of the rat in generating that plague,
and scholars still question it, a trawl through the wider extant lit-
erature about rats and the history of bubonic/pneumonic plague
suggests that these arguments are misguided. In the case of the
British Isles, the long-established popular nomenclature for black
rats as “ship” rats (as well as “house” rats) derives from their pres-
ence aboard ships, some of them carrying grain along with other
cargo, on short coastal routes. Plague could have reached the
country via infected ﬂeas traveling on humans (siehe unten) or in

4 Ruth L. Berkelsman and James M. Hughes, “The Conquest of Infectious Diseases: Who
Are We Kidding?” Annals of Internal Medicine, XIX (1993), 426–427; David T. Dennis,
“Plague in India,” British Medical Journal, CCCIX (1994), 893–894. The journal Emerging
Infectious Diseases, published from January 1995, bears witness to the existence and vigorous
growth of a scholarly community concerned with infectious diseases. It began as a quarterly
publication before becoming bi-monthly in 1999 and monthly in 2002.

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goods, but black rats themselves could not have reached Britain
without traveling by ship. Now largely displaced by the brown
or Norway rat (also known as the “wander” rat), black rats had
established a presence in Britain well before the Black Death
(siehe unten); they are still to be found in the ports and port towns
around Britain, especially in the environs and hinterlands of
London, Cardiff, and Liverpool.5

Recent archaeological evidence from London tends to sup-
port Davis’ argument for a vector other than the black rat for
plague in the fourteenth century. Although rats certainly inhabited
London and elsewhere in England both before and after the Black
Death, the mass rat deaths that are known to accompany modern
bubonic plague are not in evidence there, least of all in the
thirteenth- to fourteenth-century waterfront dumps where they
are expected to be. dennoch, an animal that prefers built struc-
tures (mainly houses and ships, as its nicknames suggest) as a habitat
is arguably more likely to die invisibly within them, leaving no
trace when those constructions are demolished. Few buildings
and ships survive from the thirteenth century, certainly not the
timber-framed, wattle and daub–walled dwellings in which most
of the human population lived. The house mice within the walls
of the British houses that have survived from the sixteenth century
onward, whether made of lath and plaster or brick, usually die

5 For the plague’s entry into England through the small port of Weymouth on the south coast
and its conveyance inland via infected goods, see Ole Benedictow, The Black Death 1346–1353: A
Complete History (Woodbridge, 2004), 126–128. According to the British Wildlife Centre, black rats
entered the country with the Romans: https://britishwildlifecentre.co.uk/planyourvisit/animals/
blackrat.html. As a possible indication of their slow spread across the country, W. R. Boelter, Der
Rat Problem (London, 1909), 8, suggests that the Norman Conquest of 1066 marked their entry,
noting that in the Wales of his day, the black rat was still known as Llgyodun Ffancon, “the French
mouse.” For the rats’ whereabouts in the ports, see the National Biodiversity Network (NBN) maps,
verfügbar unter https://species.nbnatlas.org/species/NHMSYS0000080213 (abgerufen im Juli 19, 2018); für
the import and re-export of grain into London to and from Europe in medieval England, James A.
Galloway, “One Market or Many? London and the Grain Trade of England," im gleichen (Hrsg.), Trade,
Urban Hinterland and Market Integration c. 1300–1600 (London, 2000), 23–42, 33; for the names and
habits of the black rat, https://en.wikipedia.org/wiki/Black_rat. The introduction of the brown rat
into Britain is often dated as c. 1750, but indisputable evidence places it in Europe at least from the
sixteenth century. L. Fabian Hirst, The Conquest of Plague: A Study in the Evolution of Epidemiology
(London, 1953), 123–124. Hirst (1882–1964) was bacteriologist to the municipal government of
Colombo, Ceylon (Sri Lanka), aus 1911 Zu 1915 and again from 1918 Zu 1934, having joined the
Royal Army Medical Corps during World War I, serving in Egypt and Salonica. For aspects of his
Karriere, see http://gmic.co.uk/topic/63382-serbian-re-cross-decoration-great-war/?page=2
(abgerufen im Juli 26, 2018), and William MacArthur’s foreword to Conquest, v.

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T H E U N D E R – A P P R E C I A T E D R O D E N T
inside the walls or under the ﬂoors of these homes, not in the
offen. The same pattern could hold for the house/ship rat.6

Evidence from the British government’s Indian Plague Com-
mission (1905–1907) tends to conﬁrm these observations. Both the
black rat and the grey (brown) rat were “far and away” the com-
monest rodents in early twentieth-century Bombay. The black rat,
represented in large numbers throughout the city, was described as
“essentially a house rat; it may almost be called a domesticated
Tier, living and breeding as it does in the house where people
live.” Although “typically a climbing rat,” it also burrowed into
earthen ﬂoors and walls, and its nests were often found in “little-
disturbed accumulations,” such as cotton waste, stacks of ﬁrewood,
usw., “and in recesses such as cupboards.” Furthermore, the type of
building that black rats favored typically had earthen ﬂoors and
“country tiled roofs,” were in disrepair, and contained “accumula-
tions” of rubbish and food, stores of grain, usw. Im Gegensatz, grau
(brown) rats lived mostly outside houses, “in sewers, storm water
drains, stables etc,” and nested in burrows, “even” forming extensive
burrows in the walls and ﬂoors of buildings. They never ventured
above the third ﬂoor of a house, preferring house compounds, sta-
bles, godowns (warehouses), and food and tea shops. The two spe-
cies had a common meeting ground in gullies, the lower ﬂoors of
Häuser, and warehouses.7

Housing conditions in 1900s Bombay must have closely re-
sembled those of medieval settlements, with earthen ﬂoors on
the bottom and wooden ones above and permeable walls in which
black rats could establish themselves. Bombay authorities collected
both live and dead rats, performing postmortems on more than
5,000 of the dead ones during this operation. The results were in-
conclusive. In a group of affected villages outside the city, Die
number of plague rats was “very small” in proportion to the sever-
ity of the epidemic, “notwithstanding the very thorough and ex-
tensive search made.” This experience, as Lamb noted, “points to
the danger of concluding that plague-rats are absent from an in-
fected locality unless a very thorough search is carried out.” Given
these Indian observations, and the nature of fourteenth-century

6 Barney Sloane, The Black Death in London (Stroud, 2011), 183–184.
7 George Lamb, The Etiology and Epidemiology of Plague: A Summary of the Work of the Plague
Commission (Calcutta, 1908), 9, 10.

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house construction, the lack of evidence regarding mass rat deaths,
even from the city of London, is hardly surprising.8

Recent historical scholarship indicates that London’s inordi-
nately high death rates from 1348 Zu 1353 were due to the pneu-
monic rather than the bubonic form of the disease. Advanced
bubonic plague can spread to the lungs, causing pneumonic
plague, and aerosolized droplets from persons with either form
of plague can infect others. Pneumonic plague has a rapid incuba-
tion period: Death can occur within four days; untreated, it is usu-
ally fatal. The dramatic death rates in the City of London receive
conﬁrmation in the wills of wealthy citizens who died at this time,
indicating a mortality rate of 55 Zu 60 Prozent. Material taken from
London’s East Smithﬁeld burial ground in 2011 allowed re-
searchers to sequence the Yersinia pestis genome, thus conﬁrming
the identity of plague as the cause of this epidemic and indicating
that the strain that caused the Black Death was the ancestor of
most modern strains of the disease. Crossrail excavations in 2013
London enabled a further archaeological investigation in Charter-
house Square, which revealed a plague cemetery containing a large
number of skeletons, neatly buried in layers. Dental pulp exam-
ined with the latest ancient DNA methods again revealed Yersinia
pestis as cause of death and testiﬁed to the pre-existing poor health
status of the victims, which would have rendered them highly
vulnerable to severe infection.9

Although this evidence lends substance to the role of human-
to-human infection, it diminishes, though it does not totally
undermine, the traditional role of the rat. Sources from the early
twentieth century, when plague was still a serious concern for
British colonial administrators, show that black rats were present

Ebenda., 15, 18.

8
9 For information about plague from the World Health Organization (WHO), see www.
who.int/csr/disease/plague/en/ (accessed April 1, 2019); for the London wills, Sloane, Black
Death, 103–110; for information about pneumonic plague, https://www.sciencedirect.com/
topics/pharmacology-toxicology-and-pharmaceutical-science/pneumonic-plague; “New
Findings Rewrite the Story of the Back Death,” 30 Marsch 2014, available at http://www.
channel4.com/info/press/news/new-ﬁndings-rewrite-the-story-of-the-Black-Death (ac-
cessed July 19, 2018). Kirsten I. Boc et al, “A Draft Genome of Yersinia Pestis from Victims
of the Black Death,” Nature, 478 (Oktober 27, 2011), 506–510. These ﬁndings conﬁrm the
conclusions reached by DeWitte and Slavin in “Between Famine and Death,” using earlier
ﬁndings from London material. The study of ancient DNA, pioneered in 1984, has advanced
remarkably during the last decade.

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| 177

in medieval Britain around the time of the Black Death, that they
had been present in Europe since prehistoric times, and that hu-
man individuals can act as plague transmitters independently of
black rats. Among other things, the Indian Plague Commission
found that plague was “usually conveyed from place to place by
imported rat ﬂeas, which are carried by people on their persons
or in their baggage. The human agent himself not infrequently escapes
infection” (my italics). The upshot is that the Black Death may have
reached Britain without the agency of rats in yet another way.10
Hirst—whose ﬁrst-hand studies of plague as a microbiologist
in Ceylon (modern Sri Lanka) zwischen 1912 Und 1934 gave him
an eclectic knowledge of the history of his subject—contended,
“The view that rats were scarce in the British Isles during the
fourteenth century is untenable.” Evidence produced in support
of this statement included a quotation from Geoffrey Chaucer‘s
fourteenth-century “Pardoner’s Tale,” in which a young rioter
bought poison from an apothecary “that he might his ratouns
quell.” It also contains the observation that the Old High and
Low German languages distinguished rats from mice, wohingegen
“the Aryan group of languages [gebraucht] the same word . . .
für
both.” Hirst further noted that Ælfric (C. 955–c. 1010), ﬁrst Abbot
of the Benedictine monastery at Eynsham near Oxford, used the
word “raturus,” and a ﬁfteenth-century Irish bishop is on record as
having had his library destroyed by “majores mures more vulgarly
called rati” (“large mice more vulgarly called rats”).11

Hirst also found more concrete evidence of rats. He noted
that “the record of the rocks” (fossil specimens) suggested that
“a species of rodent closely resembling Rattus rattus” had existed
in Europe since prehistoric times. He referred speciﬁcally to spec-
imens found “near Geneva, in Lombardy, Bohemia and in Crete,”
as well as the “half-digested remains” of R. rattus alexandrinianus
among the mummiﬁed remains of sacred birds in Egypt. Also to
the point is Hirst’s observation that “the free-wandering species of
ﬂea” common to the black rat is “far more likely to attack man
when it deserts the cooling body of a plague rat” than is the
nest-loving species generally found on the brown rat.12

10 Lamb, Etiology of Plague, 93.
11 Hirst, Conquest of Plague, 122.
12

Ebenda., 126, 124.

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Sources from the late twentieth and twenty-ﬁrst centuries can
also provide clues to what might have happened in past bubonic
plague outbreaks. An example from modern Madagascar, Wo
plague is now endemic in the local rodent population, illustrates
the ease with which transmission from human to human occurs,
as well as the role that small rodents other than rats play in main-
taining reservoirs of infection. In Madagascar, the rural shrew pop-
ulation, which still carries the infection without being affected by
Es, lives alongside black rats in the highlands and shares their ﬂea
Spezies. Rats may also develop resistance to the plague bacillus,
adding a further layer of complexity to the processes of infection
and making plague impossible to eradicate. Until recently, plague
infection was endemic in the island only on land over 800 meters
above sea-level. In 2017, an individual infected with bubonic
plague traveling by bus from the highlands to Antananarivo, Die
island capital, and then to the coastal city of Tamatave transmitted
pneumonic plague to thirty-one of his fellow travelers, four of
whom died, despite modern antibiotic treatment.13

As the Madagascar case demonstrates, and early twentieth-
century observers conﬁrm, rats are not needed for an outbreak
of pneumonic plague; human travelers will do. What made the
Indian epidemic of 1994 so “unexpected and dramatic” was not
so much the outbreak itself as the form that it took. Both bubonic
and pneumonic plague had already caused outbreaks in Myanmar,
Vietnam, Tanzania, Zaire, Peru, and Madagascar in the 1990s, Aber
none of them had aroused much international attention. Der 1994
Indian epidemic, Jedoch, was different. Large numbers of dead
rats and new cases of bubonic plague emerged in the Seed district
of Maharashtra state, 300 miles east of Bombay, in September. Von
September 26, ﬁfteen villages had suspect cases. Four days earlier,
Surat, a port city in Gujarat state 300 miles north of Bombay—
home to many migrant workers among its population of 1 Million
people—had reported pneumonic plague but no bubonic plague
and no unusual rat deaths. This sequence was thought to result
from epizootic plague in wild rodents spilling into the commensal

13 Pascal Boisier et al., “Epidemiologic Features of Four Successive Annual Outbreaks of
Bubonic Plague in Mahajanga, Madagascar,” Emerging Infectious Diseases, VIII (2007), 311–315;
Suzanne Chauteau et al., “Plague, a Re-Emerging Disease in Madagascar,” ebenda., IV (1998),
102. For plague in 2017 Madagascar, siehe https://www.livescience.com/60715-plague-out-
break-madagascar.html (abgerufen im Juli 10, 2019).

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| 179

rodent population in Maharashtra, resulting in primary bubonic
cases in humans with secondary pneumonic cases in workers re-
turning to Surat. Es ist, wieder, suggestive of what might have hap-
pened in British ports in 1348 if uninfected plague-ﬂea carriers, oder
incubating pneumonic cases, or even bubonic cases, had arrived on
ship at the quaysides.14

Pneumonic plague normally requires close contact with a
person already ill, but it also spreads through close contact with,
or the eating of, sick animals, not necessarily rats. In the last, spas-
modic, outbreak of plague in Britain, which occurred in Suffolk
zwischen 1906 Und 1918, two initial cases derived from rabbits,
one pneumonic (1906) and one bubonic (1911). Infected animal
species identiﬁed around the affected parish of Freston in 1906
were rat, rabbit, hare, cat, and ferret. Im 1906 outbreak, Die
affected family had eaten a rabbit brought home by the house-
holder for dinner shortly before the index case began; Die 1911
victim had cut up a rabbit, slightly injuring himself in the process,
before his illness developed.15

Herbert Bulstrode, the senior investigating ofﬁcer for the
1910 outbreak, thought that this infection had most likely arrived
with rats from the grain ships that brought cargo into the ports on
the River Orwell from San Francisco, San Nicolas, Valparaiso,
Rosario, and Alexandria. Van Zwanenberg, a twentieth-century
medical commentator, Jedoch, thought it more likely to be syl-
vatic plague, noting that the disease had spread from China and
Hong Kong in 1894 to many parts of the world, einschließlich der
Scottish city of Glasgow in 1900 (where it killed sixteen people).
In his view, it could have been an enzootic in the Suffolk area,
only accidentally infecting individuals. Citing Hirst, er hat das bemerkt
sylvatic plague tended to be pneumonic, not bubonic. Reports of
an “undue” mortality among rats at the time of the 1910 outbreak
beim Menschen, supported by subsequent local and bacteriological in-
vestigations, suggested that sylvatic plague had indeed established
itself in the two rural districts of Samford and Woodbridge, welche
lay on either side of the Orwell River in Suffolk, and close to the
Suffolk coast and the ports of Harwich and Felixstowe. Diese

14 Lamb, Etiology of Plague, 93; Dennis, “Plague in India,” 893–894.
15 For how pneumonic plague spreads, siehe https://www.livescience.com/60715-plague-
outbreak-madagascar.html (abgerufen im Juli 10, 2019). David van Zwanenberg, “The Last Epidemic
of Plague in England? Suffolk 1906–1918,” Medical History, XIV (1970), 62–74.

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| AN N E HA RD Y

180
Fälle, which demonstrated that plague infection was “fairly wide-
spread” among rats and other rodents locally, triggered a vigorous
campaign that killed more than 6,000 rats in the affected areas.16
The two histories from modern-day Madagascar and early
twentieth-century Britain witness not just to the extreme infec-
tiousness and fatality of pneumonic plague but also to the potential
role of a wider range of animal carriers than the black rat. In crowded
fourteenth-century ports, the number of rats sufﬁcient to start a train
of pneumonic infection can be small. Van Zwanenberg suggested
that the Suffolk outbreaks might constitute the model for the
behavior of plague in rural England, explaining the persisting in-
fection in the country between 1350 and the late seventeenth cen-
tury, which can be traced in burial registers and bills of mortality.
The infection would have been impossible to detect if it had run
its course as it did in Suffolk from 1906 Zu 1918; not until the
opening of a bacteriological laboratory at Ipswich Hospital in
1910 were researchers able to diagnose this twentieth-century
manifestation. Van Zwanenberg noted that conditions in early
twentieth-century Suffolk did not differ much from those of the
seventeenth century and that the Suffolk cases proved brown rats
to be as capable of carrying the infection as black rats.17

The ability of plague to become established as an endemic in-
fection in rodents and animals other than rats re-opens the ques-
tion of endemic plague reservoirs, which has been in play since
Twigg applied biological science to the role of the black rat in
1984. A recent scientiﬁc analysis that gives evidence of climate-
driven re-introductions of the bacillus providing the dynamic of
fresh epidemics does not ﬁnd support for the existence of perma-
nent plague reservoirs in medieval Europe. Noch, Carmichael dem-
onstrated the presence of a rodent reservoir of plague among Alpine
marmots in the sixteenth century. Außerdem, the evidence from

16 Zwanenberg, “Last Epidemic,” 71,102. Useful details of the local circumstances are
found in H. P. Sleigh, “Four Cases of Pneumonic Plague,” British Medical Journal, 12 Nov.
1910, 1489–1490; Herbert H. Braun, “The Recent Cases of Plague in Suffolk,” ebenda., 1490.
Anon, “Plague in Glasgow,” The Lancet, 22 Sept. 1900, 897–999. For the spread of plague in
the 1890s, see Myron Echenberg, Plague Ports: The Global Urban Impact of Bubonic Plague, 1894–
1901 (New York, 2007). Hirst, Conquest of Plague, 192, 195–196; Sleigh, “Four Cases of Pneu-
monic Plague,” 1489; Chief Medical Ofﬁcer’s Annual Report, British Parliamentary Papers,
XXXII (1911), 14–55.
17 Zwanenberg, “Last Epidemic,” 71–72.

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| 181

T H E U N D E R – A P P R E C I A T E D R O D E N T
early twentieth-century Suffolk—together with that of the modern
Vereinigte Staaten; Hirst’s descriptions of plague reservoirs among
Russian marmots and ground squirrels, South African gerbils, Und
Argentinian cavies; and the endemic infection in shrews in
present-day Madagascar—suggest that sylvatic reservoirs persisted
in Europe as elsewhere for as long as climate and ecological condi-
tions permitted. If plague entered medieval and later settlements by
human agency from sylvatic reservoirs, it would have left no trace of
mass rat deaths, as the modern case of Madagascar illustrates. Neither
Pepys nor Defoe, both acute observers, mention rat die-offs in their
respective accounts of the epidemics of 1665/6.18

The modern-day histories above witness not only to the ex-
treme infectiousness and fatality of pneumonic plague but also to
the potential role of a wider range of animal carriers than the black
rat. The indications that the rat was not widely present in
fourteenth-century English towns are not just archaeological. Al-
though the legend of the pied piper of Hamelin in Lower Saxony
dates back to c. 1300 and evidence attests to the presence of a rat
catcher there at that time, the Oxford English Dictionary traces the
ﬁrst use of the English word rat to 1592. If this dating is accurate,
urban rats did not exist in sufﬁcient numbers to constitute a serious
nuisance for many years, perhaps adequately controlled by ordinary
citizens using poison, as the reference to Chaucer above suggests.
Other creatures, such as rabbits, could also have been involved in
the diffusion of plague, as in Suffolk. The Romans introduced
rabbits into Britain as a food source, which they were to remain
as a staple in English towns and cities well into the twentieth cen-
tury. By the twelfth century, domesticated rabbits had also become
popular among the well-to-do for their fur.19

18 Graham Twigg, The Black Death: A Biological Reappraisal (London, 1984); Boris V.
Schmid et al., “Climate-Driven Introduction of the Black Death and Successive Plague Re-
introductions in Europe,” Proceedings of the National Academy of Sciences of the United States of
Amerika, CXII (2015), 3020–3025; Ann G. Carmichael, “Plague Persistence in Western
Europa: A Hypothesis,” in Green (Hrsg.), The Medieval Globe (2014), ICH, 157–191; Hirst, Conquest,
189–204; Robert Latham and William Matthews (Hrsg.), The Diary of Samuel Pepys (London,
1983), XI, 227–228; Daniel Defoe, A Journal of the Plague Year (London, 1966). Defoe, Wie-
immer, notes the killing of cats, dogs, and rats in London, in the belief that their fur carried the
poison of plague. Defoe also records that the disease entered London from the west and north,
not from the river (137, 146). I am grateful to Mary Dobson for these observations.
19 According to the Concordance of Shakespeare’s Works (primarily, 1589–1613) (http://
www.opensourceshakespeare.org/concordance/), the word rat occurs ten times therein, “rats”
eight times, ratsbane (arsenic) three times, and rat-catcher once. For domesticated rabbits, sehen

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182

| AN N E HA RD Y

The United States offers a modern example of the establishment
of a wildlife reservoir of plague—one that has survived for at least a
Jahrhundert, involving several species of rodent. Plague was ﬁrst intro-
duced into America via the Chinese population of San Francisco
on March 6, 1900. That same year, quarantine stations in Port
Townsend and New York diagnosed several cases, though the dis-
ease did not progress further into those cities. Von 1910, the United
States had three areas of known plague infestation, two on the
California seaboard and one in Washington state, as well as a small
patch on the shoreline border between Oregon and Washington.
Von 1935, the California infection had considerably extended its
range, reaching, in a broken line, from the southeastern corner
to the northwestern border just into Oregon. Plague also reached
an area in northeastern Idaho, along the border with Montana. Von
1939, the disease had marched solidly into Montana, Wyoming,
Utah, Nevada, Arizona, and New Mexico in a patchwork pattern.
Although rats appeared to be the principal agents, Kalifornien
discovered infected ground squirrels in 1924/5.20

In 1950, plague was identiﬁed in cottontail rabbits in Lea
County, New Mexico. A wildlife survey detected numerous dead
cottontail rabbits and pack rats; two specimens of each rodent had
plague bacteria in their tissue and ﬂeas. Fleas from other pack rats,
as well as from pack-rat nests and grasshopper mice were also
found to be infected. Since cottontail rabbits had never been im-
plicated in an epizootic before, health ofﬁcers in counties with
known active plague foci warned hunters of the danger involved
in handling and cleaning rabbits. Bis zum Jahr 2000, rodent reser-
voirs of bubonic plague in the western United States were present
in marmots, rabbits, kangaroo rats, deer mice, and ground squir-
rels, each carrying a different ﬂea species, none of which were
Xenopsylla cheopsis, the classic plague ﬂea, or Pulex irritans, Die

https://www.justrabbits.com/rabbit-history (abgerufen im Juli 20, 2018); Gilda O’Neill, My East
End: Memories of Life in Cockney London (London, 2000), 209. For earlier evidence, see Medical
Ofﬁcer of Health Annual Report (Birmingham, 1880), 21; ibid. (1887), 47; for cellar-kept rabbits
in eighteenth-century London, Tobias Smollett, Humphrey Clinker (New York, 1967; orig.
Kneipe. London, 1771), 153.
20 The outbreak in San Francisco has been well studied. Sehen, Zum Beispiel, Susan Craddock,
City of Plagues: Krankheit, Poverty and Deviance in San Francisco (Minneapolis, 2000); Guenter B.
Risse, Plague, Fear, and Politics in San Francisco’s Chinatown (Baltimore, 2012). Brock C.
Hampton, “Plague in the United States,” Public Health Reports, LV (1940), 1145–1149, inkl-
ing Figure 1 (1147).

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T H E U N D E R – A P P R E C I A T E D R O D E N T
human ﬂea. Human plague cases still occur in the western states, al-
though the infection has, for presumed ecological or climactic reasons,
progressed no further east. As an indicator of the ongoing danger of
infection, the Centers for Disease Control offers several information
packages on its website, including fact sheets in English and Spanish.21
The ability of rats to travel the globe by ship acquired conﬁr-
mation from at least two important sources. Following the outbreak
of plague in India c. 1894, Sir Richard Thorne Thorne, Britain’s
chief medical ofﬁcer, introduced an annual review of foreign
manifestations of plague and of its occasional appearances onboard
British ships and in British ports. On the other side of the Atlantic,
where plague was a newly established epizootic, matters were even
more serious. Link, Senior Surgeon for the Communicable Disease
Center in San Francisco, published “Plague on the High Seas,” a
special study of the connexions between plague, rats, and shipping,
In 1951. His report expressed surprise that the importance of the rat
and its ﬂea had “not been noticed until the twentieth century,” cit-
ing several literary references connecting rats and plague—from the
Bible through the works of Poseidonius (C. 135 B.C.E–c. 51 B.C.E.),
the eighteenth-century Chinese poet Shih Taonan, and the Chinese
ofﬁcial who offered a “10-cash piece” for every dead rat brought to
him during the 1894 Hong Kong epidemic. For the current (Dritte)
pandemic, Link observed that the ﬁrst recorded cases of ship-borne
plague had occurred in June 1894 on two vessels traveling from
Hong Kong to Japan and Singapore. No ship-borne cases were re-
ported in 1895, but from 1896 Zu 1938, plague was reported aboard
ships every year—332 cases in which “human, rodent, or both types
of plague were conﬁrmed by clinical or laboratory determinations.”
These cases “involved many ports of departure and arrival in 55
countries in all continents.”22

21 Link, “Plague Epizootic in Cottontail Rabbits,” Public Health Reports, XCV (1950), 696;
Alfonso Ruiz, “Plague in the Americas,” Emerging Infectious Diseases, VII (2001), 539–540. Für
a recent update, see Kiersten J. Kugler et al., “Epidemiology of Human Plague in the United
Zustände, 1900–2012,” ibid., XXI (2015), 16–22. For information about plague from the Centers
for Disease Control, siehe https://www.cdc.gov/plague/index.html (accessed April 1, 2019).
See also Arnold F. Kaufmann, John M. Boyce, and William J. Martone, “Trends in Human
Plagues in the United States,” Journal of Infectious Diseases, CXLI (1980), 522–524; Maria
Danforth et al., “Investigation of and Responses to 2 Plague Cases: Yosemite National Park,
Kalifornien. USA, 2015,” ebenda., XXII (2016), 2045–2053.
Sehen, Zum Beispiel, Medical Ofﬁcer’s Annual Report to the Local Government Board,
22
British Parliamentary Papers, XXXII (1911), 46, 123–173; Bruce C. Lowe, “The Progress and

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184

| AN N E HA RD Y
Link further described the international agreements (1903–
1944), fumigation methods, and techniques of investigation that
had attempted to end these plague-export routes, noting that only
after the development of practicable methods for rat-prooﬁng
ships in 1924/5, based on those already used in the land-based
building industry, was success ﬁnally achieved. The proof of this
success was the complete absence of any reports of plague aboard
any ship in the world since 1938. Link’s survey provides solid
evidence of the frequent, if not routine, presence of rats on ships
Vor 1925; it is difﬁcult to imagine that rats were not regular
travelers on medieval and later vessels.23

This brief longue durée perspective on the involvement of rats and
other rodents in plague history reﬂects the interdisciplinary nature
of bubonic-plague studies. On the scientiﬁc side, Hirst’s attention
to the historical traces of black rats and current rodent plague reser-
voirs in the mid-twentieth century continued in Van Zwanenberg’s
account of bubonic plague in Suffolk from 1906 Zu 1918 und in
Stenseth and his colleagues’ turn to modern biology and mathemat-
ical modeling to suggest historical patterns of bubonic-plague
Bewegungen. The Stenseth group’s publications demonstrate how
far modern science has come in its ability to make sophisticated
contributions to our understanding of the past. Archaeologists’ dis-
coveries of plague burial grounds in London and their use of
modern genome sequencing to unlock the secrets of ancient DNA
and determine the microbial cause of the Black Death have proved
invaluable in resolving alternative suggestions regarding causality, als
well as lending a stark immediacy to the human consequences. Der
assiduousness with which past generations of public-health physi-
cians recorded their work to advance medical understandings and
achievements—as evidenced by the Indian Plague Commissioners,
the early twentieth-century general practitioners in Suffolk and their
colleagues at the Local Government Board, and Link—is not to be
underestimated.

Historians have also made vast contributions to this inter-
disciplinary recovery of the rat and bubonic plague: Witness the

Diffusions of Plague through the World during 1910,” ibid., 205–255; Link, “Plague on the
High Seas,” Public Health Reports, LCVI (1951), 1467, 1468. Die Zahlen, which derive from
the Public Health Reports, IX–LCIII (1894–1938), apply to American shipping only.
23 Link, “Plague on the High Seas,” 1469–1470, 1471.

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| 185

works of Hirst and Link, who were both scientists and amateur his-
torians. Sloane’s use of the documentary evidence, particularly wills,
to evaluate the shockingly high death rates from London’s plague of
1347 brings that epidemic crisis into sharp focus. Carmichael’s un-
covering of the marmot reservoir of bubonic plague in the Alps
provides a clear-cut testimonial to Hirst’s account of reservoirs
among marmots and other rodents in various parts of the world.
This short history of the rat and the wider rodent family in
relation to bubonic plague suggests multiple ways in which differ-
ent research disciplines can contribute to the understanding of
Mortalität, morbidity, and epidemics in the past. Demographic ap-
proaches to the history of such phenomena can clarify long-term
trends in, and disruptions to, patterns of mortality, especially in
tandem with expertise in the epidemiology and symptoms of
speciﬁc disease entities. The study of recent and current disease
behaviors since 1850—using period and contemporary materials
combined with modern scientiﬁc and epidemiological methods
whenever possible—can lend insights into past disease behaviors.
Psychologie, as Lebrun’s Les hommes et la mort clearly shows, is also
pertinent to aspects of this history. Archaeological discoveries and
the still-developing technology of ancient DNA analysis can be ex-
tremely helpful; the most recent studies of London’s burial pits are
a case in point. Darüber hinaus, primary historical research involving,
sagen, the study of wills and other documentation, are essential to
creating an accurate picture of historical circumstances. Without
the collaboration of interdisciplinary methods, as the example of
the black rat and bubonic plague shows, our understanding would
surely suffer. The history of plague and the Black Death encom-
passes far more than the involvement of rats, but the more or less
enduring sylvatic reservoirs of plague infection that the rats and
their many rodent cousins constituted in the past, and still do in
the present, should not be blithely discounted.

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3 Zeitschrift für interdisziplinäre Geschichte, L:2 (Herbst, 2019), 171–185. Bild

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