Transmission of Zoonotic Diseases in the Daily Life of Ancient Pompeii and Herculaneum (79 CE, Italy): A Review of Animal–Human–Environment Interactions through Biological, Historical and Archaeological Sources

Abstract

There is no doubt that the cultural and urban environments contributed to the animal–human interaction in the daily life of the ancient Roman world. The singularity of the circumstances of the burial of Pompeii and Herculaneum, together with literary sources and the extraordinary state of preservation of the archaeological and biological material found, has provided researchers with an opportunity, unique in its kind, to reconstruct the life and ways of living of its inhabitants. This study illustrates the main drivers and mechanisms for the distribution and transmission of zoonotic diseases in these ancient Roman populations, such as (i) the large number and role that different animal species played in the ancient Roman world; (ii) the environmental conditions for the survival of parasites, pathogens and vectors; (iii) the great variety and intensity of commercial activities and occupations that presented certain risks of infections; (iv) the absence of adequate safety controls during processing, distribution and preservation of foodstuffs in unsuitable environments and some culinary habits; (v) the inadequate mechanisms of the disposal of human waste and the biotic contamination of watercourses and reservoirs; and finally (vi) the use of animals related to religious and cultural practices.

Full text

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Citation: Tanga, C.; Remigio, M.;
Viciano, J. Transmission of Zoonotic
Diseases in the Daily Life of Ancient
Pompeii and Herculaneum (79 CE,
Italy): A Review of
Animal–Human–Environment
Interactions through Biological,
Historical and Archaeological
Sources. Animals 2022,12, 213.
https://doi.org/10.3390/
ani12020213
Academic Editor: Nicole
Gottdenker
Received: 28 November 2021
Accepted: 9 January 2022
Published: 17 January 2022
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animals
Review
Transmission of Zoonotic Diseases in the Daily Life of Ancient
Pompeii and Herculaneum (79 CE, Italy): A Review of
Animal–Human–Environment Interactions through Biological,
Historical and Archaeological Sources
Carmen Tanga 1, Marta Remigio 2and Joan Viciano 3,*
1Department of Legal Medicine, Toxicology and Physical Anthropology, Faculty of Medicine, University of
Granada, Avenida de la Investigación 11, 18071 Granada, Spain; carmentanga@correo.ugr.es
2Independent Researcher, Strada Fonte Borea 1, 65125 Pescara, Italy; martaire3@gmail.com
3Department of Medicine and Ageing Sciences, ‘G. d’Annunzio’ University of Chieti-Pescara, Via L. Polacchi
11-13, 66100 Chieti, Italy
*Correspondence: joan.viciano@unich.it
Simple Summary:
There is a wide range of historical, archaeological and biological sources to help
understand the nature of health and disease in the ancient Roman world. In this study, we examine
various predisposing elements typical of the urban environment in relation to zoonotic diseases in
the ancient Roman cities of Pompeii and Herculaneum, both devastated by pyroclastic surges as a
consequence of the eruption of Mount Somma–Vesuvius in 79 CE. We analyse the meaning and value
of the sources to develop an understanding of the many features of everyday life in these cities. We
make judgements about issues relevant to the reconstruction of the past, and finally, we synthesize
evidence from different sources to construct historical explanations and arguments in relation to the
transmission of zoonotic diseases in these ancient Roman populations.
Abstract:
There is no doubt that the cultural and urban environments contributed to the animal–
human interaction in the daily life of the ancient Roman world. The singularity of the circumstances
of the burial of Pompeii and Herculaneum, together with literary sources and the extraordinary state
of preservation of the archaeological and biological material found, has provided researchers with
an opportunity, unique in its kind, to reconstruct the life and ways of living of its inhabitants. This
study illustrates the main drivers and mechanisms for the distribution and transmission of zoonotic
diseases in these ancient Roman populations, such as (i) the large number and role that different
animal species played in the ancient Roman world; (ii) the environmental conditions for the survival
of parasites, pathogens and vectors; (iii) the great variety and intensity of commercial activities and
occupations that presented certain risks of infections; (iv) the absence of adequate safety controls
during processing, distribution and preservation of foodstuffs in unsuitable environments and some
culinary habits; (v) the inadequate mechanisms of the disposal of human waste and the biotic
contamination of watercourses and reservoirs; and finally (vi) the use of animals related to religious
and cultural practices.
Keywords:
zoonosis; human–animal interaction; zooarchaeology; archaeology; palaeopathology;
physical anthropology; habits and lifestyle; status of health and disease; Pompeii and Herculaneum
1. Introduction
Zoonosis (from ancient Greek
ζ
Animals 2022, 12, x 2 of 39
humans in both rural and urban settings where humans live [1]. The infectious agents
involved constitute a diverse range of microbial pathogens such as bacteria, viruses,
parasites, fungi, rickettsiae and prions, with a variety of animal vectors or reservoirs,
including livestock, companion animals and wildlife [2]. Zoonotic infections result in a
wide range of diseases with great socio-economic impact on populations. The most recent
example of the worldwide socio-economic impact of zoonosis is the ongoing ‘severe acute
respiratory syndrome Coronavirus 2′ (SARS-CoV-2) pandemic. The current respiratory
illness responsible for the COVID-19 pandemic is the biggest global challenge since World
War II due to the sudden halt in economic activity in both advanced and developing
countries, including global poverty that has increased for the first time since 1990 [3]. The
reported number of deaths worldwide as of 18 November 2021 is 5.1 million [4]. Based on
epidemiological data, the earliest cases of documented SARS-CoV-2 were directly linked
to markets selling wild animals in Wuhan (China), without any epidemiological link to
other localities, including the Wuhan Institute of Virology (WIV)—the subject of
considerable speculation [5]. Although there are precedents for laboratory incidents
leading to isolated infections, including SARS-CoV [6], currently, there is no evidence that
any early cases of SARS-CoV-2 had any connection to the WIV or any other laboratory.
Thus, the most parsimonious explanation for the origin of SARS-CoV-2 is a zoonotic event
[5]. Several studies show that bat and pangolin species are a natural reservoir of SARS-
CoV-2-like coronaviruses with the potential to infect humans (e.g., [7–9]). Although these
species are considered to be the hosts from which this fatal pandemic originated,
currently, the animal reservoir for SARS-CoV-2 has not been identified.
Animals play an essential role in maintaining infection in nature and contribute in
varying degrees to the distribution and actual transmission of infection in human and
animal populations. These zoonotic infections have a variety of transmission mechanisms,
such as (i) transmission by animal bites or scratches [10,11]; (ii) zoonotic pathogens
originating from food animals can reach people through the direct faecal—oral route,
contaminated animal food products, improper food handling along the food-processing
chain or inadequate cooking [2,12–14]; (iii) vectors can transmit zoonotic diseases either
actively—by biological vectors, frequently arthropods, such as mosquitoes, ticks, fleas and
lice, that may carry pathogens that can multiply within their bodies and be delivered to
new hosts, usually by biting—or passively—by mechanical vectors, such as flying insects
and other animals such as birds and rats, that can pick up infectious agents on the outside
of their bodies and transmit them through physical contact. Even humans, for example,
those who are at increased risk of exposure to certain animals, such as livestock farmers
and veterinarians, can also unintentionally become carriers of zoonotic pathogens and
transmit them to other members of the community [15,16]; and (iv) soil and water
resources contaminated with manure may contain a wide variety of zoonotic pathogens,
representing an important risk for the transmission of zoonotic diseases [17,18].
ῷ
2. Zoonotic Diseases in Ancient Times: Biological, Archaeological and Literary Evidence
Zoonotic diseases have been in existence for thousands of years and have been
hypothesized to have afflicted ancestors of humans from the beginning of the hominin
evolution. Food resources were based on the gathering of plants and wild fruits and
hunting; therefore, exposure to animal pathogens and cross infection was likely a constant
threat to daily life [19]; for example, the analysis of fossilised skeletal remains of hominins
has revealed evidence of tuberculosis—likely of bovine origin—[20,21] and chronic
brucellosis [22,23]. Numerous lines of archaeological evidence and biological data
demonstrate that outbreaks or epidemics of zoonotic infections were most likely
experienced around 11,000–9000 BCE in southwest Asia with the ‘First Agriculture
Revolution’, when communities of sedentary foragers began to emerge, and the rearing
of domesticated livestock developed with the aim of founding a farming economy that
ν
[zôion, animal] and
ν
ó
σ
o
ς
[nósos, disease]) is defined
as any disease or infection naturally transmissible from vertebrate animals to humans
in both rural and urban settings where humans live [
1
]. The infectious agents involved
constitute a diverse range of microbial pathogens such as bacteria, viruses, parasites, fungi,
Animals 2022,12, 213. https://doi.org/10.3390/ani12020213 https://www.mdpi.com/journal/animals

Animals 2022,12, 213 2 of 37
rickettsiae and prions, with a variety of animal vectors or reservoirs, including livestock,
companion animals and wildlife [
2
]. Zoonotic infections result in a wide range of diseases
with great socio-economic impact on populations. The most recent example of the world-
wide socio-economic impact of zoonosis is the ongoing ‘severe acute respiratory syndrome
Coronavirus 2
0
(SARS-CoV-2) pandemic. The current respiratory illness responsible for the
COVID-19 pandemic is the biggest global challenge since World War II due to the sudden
halt in economic activity in both advanced and developing countries, including global
poverty that has increased for the first time since 1990 [
3
]. The reported number of deaths
worldwide as of 18 November 2021 is 5.1 million [
4
]. Based on epidemiological data, the
earliest cases of documented SARS-CoV-2 were directly linked to markets selling wild
animals in Wuhan (China), without any epidemiological link to other localities, includ-
ing the Wuhan Institute of Virology (WIV)—the subject of considerable speculation [
5
].
Although there are precedents for laboratory incidents leading to isolated infections, in-
cluding SARS-CoV [
6
], currently, there is no evidence that any early cases of SARS-CoV-2
had any connection to the WIV or any other laboratory. Thus, the most parsimonious
explanation for the origin of SARS-CoV-2 is a zoonotic event [
5
]. Several studies show that
bat and pangolin species are a natural reservoir of SARS-CoV-2-like coronaviruses with
the potential to infect humans (e.g., [
7
–
9
]). Although these species are considered to be
the hosts from which this fatal pandemic originated, currently, the animal reservoir for
SARS-CoV-2 has not been identified.
Animals play an essential role in maintaining infection in nature and contribute in
varying degrees to the distribution and actual transmission of infection in human and
animal populations. These zoonotic infections have a variety of transmission mechanisms,
such as (i) transmission by animal bites or scratches [
10
,
11
]; (ii) zoonotic pathogens originat-
ing from food animals can reach people through the direct faecal—oral route, contaminated
animal food products, improper food handling along the food-processing chain or inad-
equate cooking [
2
,
12
–
14
]; (iii) vectors can transmit zoonotic diseases either actively—by
biological vectors, frequently arthropods, such as mosquitoes, ticks, fleas and lice, that
may carry pathogens that can multiply within their bodies and be delivered to new hosts,
usually by biting—or passively—by mechanical vectors, such as flying insects and other
animals such as birds and rats, that can pick up infectious agents on the outside of their
bodies and transmit them through physical contact. Even humans, for example, those who
are at increased risk of exposure to certain animals, such as livestock farmers and veterinar-
ians, can also unintentionally become carriers of zoonotic pathogens and transmit them to
other members of the community [
15
,
16
]; and (iv) soil and water resources contaminated
with manure may contain a wide variety of zoonotic pathogens, representing an important
risk for the transmission of zoonotic diseases [17,18].
2. Zoonotic Diseases in Ancient Times: Biological, Archaeological and
Literary Evidence
Zoonotic diseases have been in existence for thousands of years and have been hypoth-
esized to have afflicted ancestors of humans from the beginning of the hominin evolution.
Food resources were based on the gathering of plants and wild fruits and hunting; there-
fore, exposure to animal pathogens and cross infection was likely a constant threat to daily
life [
19
]; for example, the analysis of fossilised skeletal remains of hominins has revealed
evidence of tuberculosis—likely of bovine origin—[
20
,
21
] and chronic brucellosis [
22
,
23
].
Numerous lines of archaeological evidence and biological data demonstrate that outbreaks
or epidemics of zoonotic infections were most likely experienced around 11,000–9000 BCE
in southwest Asia with the ‘First Agriculture Revolution’, when communities of sedentary
foragers began to emerge, and the rearing of domesticated livestock developed with the
aim of founding a farming economy that produced food rather than collecting it from
the wild [
24
]. The first animals to be domesticated are thought to have been sheep (ca.
11,000 BCE) and goats (ca. 8000 BCE). Both animals were used for their meat, milk and

Animals 2022,12, 213 3 of 37
coats and became the main system of food production and for the elaboration of a wide
range of secondary products [25–32].
On the other hand, written testimonies sometimes contain mention of human and
animal sufferings. The oldest of the veterinary documents is the Kahun or el-Lahun papyrus
(ca. 2230–1800 BCE), kept in the University College Museum of London. It deals with the
diseases of livestock and fish, without reference to human diseases [
33
,
34
]. By contrast,
the Ebers papyrus (ca. 1500 BCE) mentions human diseases [
35
]. Both documents, corre-
sponding to the Middle Kingdom of Egypt, show that it is one thing to know diseases and
another, very different thing, to be aware of the reciprocal animal–human or human–animal
contagion. Among other reasons, this was due to their ignorance of pathogen agents in
nature, although some hygienic standards were born from simple observation (e.g., rejec-
tion of corpses, filth, etc.). As an example, historical references to the bubonic plague are
found in the Iliad of Homer (ca. 1000 BCE), as well as in the History of the Peloponnesian War
of Thucydides (430–426 BCE), in the Bible in the Book of Exodus, and in the Metamorphoses of
Ovid (8 CE); references to anthrax are also found in the Georgics of Virgil (29 BCE), in De
Medicina of Celsus (ca. 47 CE), in the Naturalis Historia of Pliny the Elder (77 CE) and in De
Differentiis Febrium of Galen (ca. 200 CE) [36–40].
Today we know enough about several pathogen agents and their host spectrum that
we can interpret some endemic situations and epidemics, analysing the ancient written
testimonies together with archaeological evidence and biological data. What is indisputable
is that the environmental, physical and human factors that affect the onset of disease are
numerous and variable in geographical, historical and social terms. In classical Rome
or Athens, Constantinople, pre-industrial London and many other examples, there were
both predisposing and determining factors that made the urban environment ideal for
the emergence of an epidemic or the maintenance of an endemic. In this study, we will
examine various predisposing elements typical of the urban environment in relation to
zoonotic diseases in the ancient Roman cities of Pompeii and Herculaneum, both devastated
by pyroclastic surges as a consequence of the eruption of Mount Somma–Vesuvius in 79 CE.
3. Historical Background of the Roman Cities of Pompeii and Herculaneum
3.1. Geographical Setting
The ancient cities of Pompeii and Herculaneum were in the Gulf of Naples (region of
Campania, southern Italy) (Figure 1). This region was a long volcanic plateau that spread
from the Volturno river in the north to the Sarno river in the south, and it was divided into
two areas by Mount Somma–Vesuvius. Pompeii was on the southeastern side of Mount
Somma–Vesuvius, on a volcanic spur up to 40 m above the sea level overlooking the
Sarno river. Herculaneum, on the western side, spread along the low hillsides of Mount
Somma–Vesuvius, which ended in a cliff overlooking the ancient coastline [
41
,
42
]. Pompeii
was a commercial town of strategic importance because it lay on the only route linking
the north and south and connected the seaside area with the inland fertile agricultural
region. The urban centre (forum) was home to the city’s main civic, commercial and
religious institutions. Due to its views and moderating maritime influence, Herculaneum
was considered an ideal resort and residential town, with fewer public and commercial
buildings than Pompeii [43–45].
3.2. The Eruption of Mount Somma–Vesuvius in 79 CE
The fame of Pompeii and Herculaneum is due to the catastrophic volcanic eruption,
whose dating was recently re-assessed to October of 79 CE based on new epigraphic testi-
monies, archaeological finds and volcanological data [
46
–
48
]. The details of the sequence
of relevant disastrous events consequent to the Vesuvian eruption have been handed down
to us by Pliny the Younger, the nephew of Pliny the Elder—an important Roman writer
and naturalist, as well as the admiral of the Roman fleet. In the form of two letters that he
wrote to Tacitus—a Roman historian—to inform him of his uncle’s death, Pliny the Younger
gave an eyewitness account of the eruption in a detailed, graphic and objective way. The

Animals 2022,12, 213 4 of 37
sequence of events described by Pliny the Younger and the volcanological
evidence [49–51]
has allowed two main phases of the eruption to be established. The first phase was charac-
terised by a widespread dispersal of a cloud of ash, pumice and gases from a high eruptive
column, and the second phase was characterised by six pyroclastic surges, which caused
extensive destruction in a portion of territories of the Campania region, burying the entire
cities of Pompeii and Herculaneum under tens of metres of pyroclastic material as well as
most of the other towns and aristocratic villas surrounding the volcano, such as Oplontis,
Stabiae and Boscoreale [
42
]. In Pompeii, the major casualties during the first phase resulted
from roof collapses under the weight of the pyroclastic material that rained down during
the eruption. Those who abandoned the buildings were killed by suffocation due to the
ash-rich atmosphere and harmful gases. On the contrary, in Herculaneum, the major casual-
ties were due either to thermal shock or physical trauma as a result of the kinetic energy of
the pyroclastic surges [52–54].
Animals 2022, 12, x 4 of 38
letters that he wrote to Tacitus—a Roman historian—to inform him of his uncle’s death,
Pliny the Younger gave an eyewitness account of the eruption in a detailed, graphic and
objective way. The sequence of events described by Pliny the Younger and the
volcanological evidence [49–51] has allowed two main phases of the eruption to be
established. The first phase was characterised by a widespread dispersal of a cloud of ash,
pumice and gases from a high eruptive column, and the second phase was characterised
by six pyroclastic surges, which caused extensive destruction in a portion of territories of
the Campania region, burying the entire cities of Pompeii and Herculaneum under tens of
metres of pyroclastic material as well as most of the other towns and aristocratic villas
surrounding the volcano, such as Oplontis, Stabiae and Boscoreale [42]. In Pompeii, the major
casualties during the first phase resulted from roof collapses under the weight of the
pyroclastic material that rained down during the eruption. Those who abandoned the
buildings were killed by suffocation due to the ash-rich atmosphere and harmful gases.
On the contrary, in Herculaneum, the major casualties were due either to thermal shock or
physical trauma as a result of the kinetic energy of the pyroclastic surges [52–54].
Figure 1. Map showing, with their Latin names, the main cities and towns affected by the eruption
of Mount Somma–Vesuvius in 79 CE.
3.3. Investigation and Interpretation of the Different Information Sources
The integral preservation of Pompeii and Herculaneum, achieved through rapid burial
by volcanic deposits tens of metres thick, has made it possible to reconstruct certain
aspects of daily life, thanks to the discovery of private houses, public buildings and the
skeletal remains of its inhabitants, representing an absolute unicum [55]. Buildings are
often preserved up to the second floor, as well as wooden furniture, mosaics, frescoes,
sculptures and other bronze and iron objects [46,56–60]. Even diverse organic material
such as food (e.g., bread, cheese, olives, garum, grains, lentils, beans, fresh and dried fruit
such as pomegranates, figs, walnuts, almonds) [46,61–63] and remains of human faeces
[64,65] are well-preserved. Thus, the singularity of the circumstances of burial of these
two cities, together with the exceptional nature of the cause of death of its inhabitants and
Figure 1.
Map showing, with their Latin names, the main cities and towns affected by the eruption of
Mount Somma–Vesuvius in 79 CE.
3.3. Investigation and Interpretation of the Different Information Sources
The integral preservation of Pompeii and Herculaneum, achieved through rapid burial
by volcanic deposits tens of metres thick, has made it possible to reconstruct certain
aspects of daily life, thanks to the discovery of private houses, public buildings and the
skeletal remains of its inhabitants, representing an absolute unicum [
55
]. Buildings are
often preserved up to the second floor, as well as wooden furniture, mosaics, frescoes,
sculptures and other bronze and iron objects [
46
,
56
–
60
]. Even diverse organic material
such as food (e.g., bread, cheese, olives, garum, grains, lentils, beans, fresh and dried
fruit such as pomegranates, figs, walnuts, almonds) [
46
,
61
–
63
] and remains of human
faeces [
64
,
65
] are well-preserved. Thus, the singularity of the circumstances of burial of
these two cities, together with the exceptional nature of the cause of death of its inhabitants
and the extraordinary state of preservation of the archaeological and biological material

Animals 2022,12, 213 5 of 37
found, has provided researchers with an opportunity, unique in its kind, to reconstruct the
life and ways of living of its inhabitants.
In recent decades, the palaeobiological studies carried out on the bone and dental hu-
man remains of the victims of the eruption has allowed the obtention of useful information
not only to understand the habits and lifestyle of the ancient inhabitants, but also the status
of health and diseases with which the general population was affected (e.g., [
41
,
66
–
72
]).
Although bone and dental human remains constitute the primary source through which
it is possible to derive direct information on the diseases suffered by the ancient inhabi-
tants, osteological remains from animal assemblages are no less important to provide data
that can help to elucidate the close coexistence and animal–human interaction, and the
animal-related infections transmitted to humans in the community. Therefore, to provide
an in-depth picture of zoonotic diseases of the past, scientific research should not be based
exclusively on the human and animal skeletal remains, but the interaction with other data
inferable from archaeological evidence is essential, including the urban planning and other
biological remains of organic-based nature, but without neglecting information obtainable
from literary, epigraphical and artistic sources [73,74].
4. Animal–Human–Environment Interaction in the Daily Life at Pompeii
and Herculaneum
4.1. Population Density and Housing
At the onset of the Common Era, the Mediterranean region was considered a relatively
stable epidemiological pool of diseases to which the population had adapted. However,
with its military and commerce expansion and slave trade, the Roman Empire altered this
situation through progressive contact with important groups of diseases from neighbouring
regions of the Middle and Far East, North Africa and southern and western Europa. Thus,
most of the new epidemics that devastated the Roman Empire were introduced by soldiers,
merchants, slaves or immigrants [
75
,
76
]. In fact, the expansion of the Roman Empire at-
tracted many immigrants to large cities in search of better opportunities, generating several
direct health consequences. On the one hand, immigrants constituted an immunologically
virgin mass for resident pathogens and those from origins other than their own; on the
other hand, they brought their own parasites, vectors and pathogens with them and spread
them in their new habitat [
75
,
77
–
79
]. In addition, immigration also led to the indirect
spread of health-related diseases due to overcrowding and overpopulation in cities, the
inevitable lack of hygiene in substandard housings or the systematic non-existence of
adequate waste-disposal mechanisms [80,81].
The urban population of Pompeii and Herculaneum, particularly the wealthy social class,
lived in a private single-family residence (domus), while those of the less favoured social
class lived in tenement block housing with numerous families (insulae). In the insula, a large
part of the population lived in the upper floors over their own shops (tabernae)or workshops,
in small bedrooms (cubicula), in various types of guesthouses (hospitia,stabularia,cauponae,
deversoria) or simply on the street. The insulae accommodated high population densities due
to the advantageous use of the housing as a consequence of the reduced interior spaces and
a greater number of divisions [
82
–
85
]. They were the cheapest, but the indigents and those
who could not afford accommodation when nightfall came—the wages were paid daily—
had to resort to other means. Large shanty and slum areas (tuguria) were an option [
85
,
86
].
The streets of the city were also occupied due to the generous architecture full of arches,
vaults, bridges, colonnades, stairs and terraces [
82
,
85
,
87
,
88
]. Thus, in ancient Roman cities,
areas were generated where people lived together in suffocating proximity while garbage,
faeces, urine and stagnant water accumulated, along with their associated microfauna
(e.g., helminths, protozoa, flies, mosquitoes, fleas, bedbugs, lice and other insects) and
their natural hosts (people, domestic animals, rodents, birds) [
75
,
80
–
82
]. Table 1shows the
estimated population density for the cities of Pompeii and Herculaneum compared to other
major cities of the Roman Empire (data extracted from [89]).

Animals 2022,12, 213 6 of 37
Table 1.
Total area, population density and estimated population of Pompeii and Herculaneum com-
pared with other major ancient cities of the Roman Empire.
Site Modern
Country Total Area (ha) Population Density
(People per ha)
Estimated
Population
Roma Italy 1783 518 923,406
Alexandria Egypt 972 422 410,535
Antioch Turkey 399 313 124,936
Carthage Tunisia 343 298 102,079
Athens Greece 225 429 96,429
Ephesus Turkey 263 272 71,587
Lugdunum France 170 357 60,714
Londinium
United Kingdom
160 250 40,000
Ostia Italy 154 227 35,017
Neapolis Italy 82 275 22,550
Verulamium
United Kingdom
90 183 16,500
Fregellae Italy 80 125 10,000
Pompeii Italy 60 166 9938
Emerita
Augusta Spain 81 120 9720
Volubilis Morocco 43 211 9058
Calleva
Atrebatum
United Kingdom
45 80 3600
Verona Italy 52 68 3525
Augusta
Praetoria Italy 41 83 3417
Italica Spain 49 65 3178
Iulia Valentia
Banasa Morocco 15 183 2738
Herculaneum Italy 20 115 2290
Luna Italy 23 80 1840
Conimbriga Portugal 23 66 1519
Emporiae Spain 21 63 1313
ha, hectare.
The distribution and transmission of zoonotic diseases were not only influenced by
the housing spaces but also by the different materials used in the construction of the houses.
For example, adobe, wattle or heather or cane roofs were good settlements for insects and
rodents. Even furniture could host insects and rodents, especially the beds (lectus cubicularis).
The iconography and archaeological artefacts of Pompeii and Herculaneum clearly show the
main types and uses of the bed during the Roman Empire. In Roman culture, the richest
beds had mattresses stuffed with wool or feathers, while the mattresses of the popular
classes were stuffed with straw. They generally lacked sheets but had quilts that served to
cover them made with very rich textiles, as well as wool or fur blankets [
90
,
91
]. In this way,
the beds could not only be accessible to insects and rodents, but also the bedclothes could
contribute to the biotic load. One example is carbuncle or anthrax, an infection caused by
the bacterium Bacillus anthracis. It is a zoonosis acquired by livestock (cattle, sheep, goats,
horses and pigs), with humans being the accidental host. The main risk of contracting the
infection is through contact with infected animals or their products, such as hides, fur or
wool [
36
]. The Latin poet Virgil (Georgics III.478–566; 1st century BCE) cites an epidemic of
anthrax that affected the eastern Alps and describes the symptoms in animals: “[
. . .
] then
the eyes blaze and the breath is drawn deeply, at times with heavy groans, the depths of the chest
strained by long sobs, black blood flows from the nostrils, and the coarse tongue chokes the blocked
throat.” and in humans: “[
. . .
] if anyone handled their hateful clothing, feverish blisters and foul
sweat would cover his stinking limbs, and he’d not long to wait before the accursed fire was eating
his infected body” (translated by [
92
]). Hippocrates, Ovid, Galen and Pliny the Elder also
mention anthrax-like epidemics [
93
,
94
], although a more “scientific” description is found
in De Medicina by the Roman physician Celsus [94].

Animals 2022,12, 213 7 of 37
In the domestic environment, commensal rodents such as the house mouse (Mus
musculus) live in close contact with humans and domestic animals, where man-made struc-
tures provide shelter and abundant food supplies. Thus, the house mouse depends on
humans for its food and habitat and cannot survive without human presence [
95
]. Under
commensal conditions, the house mouse is destructive to food supply and human property,
undermining and weakening wooden buildings, destroying clothing and gnawing the
furniture [
96
]. In addition to the food they eat, they also contaminate human food supplies
with their urine and excrement and can spread various zoonotic pathogens, such as bacte-
ria, viruses and helminths [
97
,
98
] through direct transmission (e.g., bite, contact), through
ectoparasites (e.g., ticks, fleas, mites and lice) or through endoparasites (e.g., nematodes,
cestodes and trematodes) that infest them [
99
]. Zooarchaeological evidence in Pompeii
registers a rise in the frequency of house mice and a concomitant decline in the wood
mouse (Apodemus sylvaticus; see Section 4.3.4. Wild animals), with which the house mouse
competes, coincident with the intensification of urbanization in the city [
100
]. Numerous
skeletal remains of Mus musculus have been found in both Pompeii and Herculaneum, in-
dicating a high density of the house mouse in these cities [
100
,
101
]. This could be one
of the reasons why the houses had domestic animals such as cats to control rodents (see
Section 4.3.3. Household animals), which could also be carriers of zoonotic diseases.
4.2. Trade Routes
The archaeological finds concerning port structures, together with literary sources,
attest that the port of Pompeii not only played an important role in the economy of the
Campania region—especially for agricultural produce—but also allowed imports to and
exports from Pompeii even from cities beyond the Empire due to its proximity to the port of
Puteoli—one of the most important ports of Rome. In addition, Pompeii was the entrepôt
for the Sarno river valley, which allowed it to trade with other inland cities through fluvial
networks. By contrast, Herculaneum had a relatively small port with an economy that served
local needs [
43
,
44
]. In this context, the Roman world represented globalization before this
term existed, establishing an extensive trade network, not only maritime and fluvial, but
also terrestrial. This allowed animals and their products, as well as people and pathogens,
to spread through the cities. The dead spaces between the cargoes carried by carriages and
ships allowed animals such as rodents to hide [
102
]. Rodents are the reservoir for many
infectious organisms, which, if transmitted to humans or domestic animal populations, can
cause disease outbreaks, often with high morbidity and some mortality.
Together with the house mouse (Mus musculus) mentioned above, the black rat (Rattus
rattus; also known as the roof rat, ship rat or house rat) is also a serious pest in urban and
rural environments [
103
]. Originating from southeast Asia, the black rat arrived in the
Mediterranean basin by two different routes: (i) terrestrial or sea-trade with Mesopotamia,
where skeletal evidence of black rats was found in Syria in 3500 BCE [
104
]; and (ii) sea-
trade across the Indian Ocean to the Red Sea ports of Egypt, where partially digested
skeletal remains were discovered in the stomach of a mummified cat and in mummified
birds of prey, some of them dated from Roman times [
105
,
106
]. With the expansion of the
Roman Empire and the increase in the human population in several cities, especially Rome,
large commercial flows were generated that probably facilitated the transport of black rats
through the different countries since the intense military and commercial activities of the
Roman Empire linked the Near East (Syria, Judea) and the north coast of Africa (Egypt,
Cyrenaica) with the western provinces [
106
]. Although there are a few black rat finds before
the Roman times in Europe—the most reliable finds come from southwestern Slovenia and
date between 1100 and 800 BCE [
107
]—the zooarchaeological records increased from the
Roman period across Europe [
108
–
111
]. In particular, zooarchaeological finds show that by
200–100 BCE, the black rat was present in Pompeii [112,113].
Although the black rats live unprotected in the wild nature of the Mediterranean,
they tend to depend on human dwellings and food storage to thrive [
102
]. Black rat
populations not only cause extensive economic damage to crops, stored food, farms,

Animals 2022,12, 213 8 of 37
industries and households, but they also harbour and spread diverse zoonotic pathogens,
such as viruses (e.g., hantavirus), bacteria (e.g., Leptospira interrogans), protozoa (e.g.,
Toxoplasma gondii) and helminths (e.g., Hymenolepis spp.) [
103
]. The dense urbanism of
the insulae, the baths (thermae), the markets (tabernae), the shops and workshops, the food
houses and the taverns (thermopolia,cauponae,popinae) could favour the establishment of
rodent populations so that the distribution of the black rat within Roman cities, including
Pompeii and Herculaneum, could follow a pattern marked by the availability of access to
shelter, food and garbage [100,102].
4.3. Livestock, Wild, Exotic and Household Animals
One of the most obvious and far-reaching differences between the life of “developed”
and mechanized countries of modern times and that of the ancient Roman world lies in the
much larger role that animals of all types played in the latter. In fact, there were few aspects
of human activity, whether at work or leisure, in which animals were not involved [114].
4.3.1. Exotic Animals
There is no doubt that great mass events were the most popular entertainment event
during the imperial era. According to the classical Roman vision of leisure, the great public
games (ludi publici) were an activity to satisfy the people’s desire for escape and recreation,
as well as to guarantee good social order by offering a palliative. Famous is the Juvenal’s
expression panem et circenses (bread and games) (Satire X.77–81; 2nd century CE), with
which the poet describes the practice of the aediles (the town authorities) of providing free
grain and costly public games as a mean to gain political power and to get the consent of the
citizens, distracting them from the social problems they were suffering. Of great importance
were the venationes, as well as the damnatio ad bestias shows in which wild and dangerous
animals participated, imported to the city expressly for this purpose. Venationes were a
form of entertainment that involved hunting and killing wild animals, while damnatio ad
bestias was a particular type of death penalty where those convicted were executed by
animals [
115
,
116
]. As the Empire expanded, exotic animals such as lions, leopards, tigers,
panthers, rhinos, crocodiles, hippos and even elephants began to be incorporated alongside
animals of local origin, such as wild boars, foxes, bulls, wolves and bears [
117
] (Figure 2a).
All these animals were captured in their place of origin, and transporting them to the
Italian peninsula required covering very long distances by land and sea. According to
the Roman writer and statesman Symmachus in his letters (Epistulae IV–VI; 5th century
BCE), many animals died during transportation, but many others reached the circus and
were used as entertainment [
114
–
116
,
118
]. Although most of their meat was destined to
feed its congeners who waited in the vivarium cages for future shows, it could be donated
to the public or to the authorities as an act of generosity from the emperor. This gesture
was very well received among the attendees, whose purchasing power was not enough
to frequently access quality protein. In any case, it was an undoubted potential route of
entry for zoonotic pathogens. However, there are no references relating to animals destined
for venationes and damnatio ad bestias shows with different diseases, whether their own or
human [81,118].
4.3.2. Livestock
Livestock was among the main resources of Roman society, offering the Mediterranean
terrain many animal species with which to raise livestock and obtain good-quality products
derived from such [
119
]. The main sources of information on Roman livestock include
ancient Latin treatises, such as those by Cato the Elder (De Agri Cultura; 2nd century BCE),
Varro (Res Rusticae; 1st century BCE), Virgil (Georgics; 1st century BCE), Columella (De Re
Rustica; 1st century CE) and Pliny the Elder (Naturalis Historia; 1st century CE). Several
artistic representations in mosaics, frescoes and sculptures testify to the implements used
in agriculture and livestock and, occasionally, metal parts of these implements have been
brought to light during the diverse archaeological excavations. The types of livestock of

Animals 2022,12, 213 9 of 37
the ancient Roman world were mainly cattle, equines, sheep, pigs, poultry, rabbits, bees
and fish.
Animals 2022, 12, x 9 of 38
4.3.2. Livestock
Livestock was among the main resources of Roman society, offering the
Mediterranean terrain many animal species with which to raise livestock and obtain good-
quality products derived from such [119]. The main sources of information on Roman
livestock include ancient Latin treatises, such as those by Cato the Elder (De Agri Cultura;
2nd century BCE), Varro (Res Rusticae; 1st century BCE), Virgil (Georgics; 1st century BCE),
Columella (De Re Rustica; 1st century CE) and Pliny the Elder (Naturalis Historia; 1st
century CE). Several artistic representations in mosaics, frescoes and sculptures testify to
the implements used in agriculture and livestock and, occasionally, metal parts of these
implements have been brought to light during the diverse archaeological excavations. The
types of livestock of the ancient Roman world were mainly cattle, equines, sheep, pigs,
poultry, rabbits, bees and fish.
Figure 2. (a) Mosaic of a lion attacking a leopard (House of the Mosaic Doves, Pompeii). Despite the
presence of vegetation in this scene, it seems likely that whoever designed this mosaic had observed
these animals not in their natural habitat, but in the arena of Pompeii’s amphitheatre. Animals were
sometimes forced to fight each other after having been starved and tormented to increase their
aggression. At other times, animals were part of “hunts” (venationes) or pitted against gladiators or
criminals condemned to die (damnatio ad bestias) [Author: MatthiasKabel. Distributed under a CC
BY-SA 2.0 license. Available online: https://cutt.ly/dRCLc99. Accessed: 2 February 2021]. (b) Plaster
cast of a pig found during the excavation in Villa Regina (Boscoreale). It certifies the breeding of pigs
on Vesuvian farms [Author: RealCarlo. Distributed under a CC BY 2.0 license. Available online:
https://cutt.ly/KRCLFFl. Accessed: 30 October 2021]. (c) Fresco of a rabbit and four figs (from
Herculaneum) [Author: Amphipolis. Distributed under a CC BY-SA 2.0 licence. Available online:
Figure 2.
(
a
) Mosaic of a lion attacking a leopard (House of the Mosaic Doves,Pompeii). Despite the
presence of vegetation in this scene, it seems likely that whoever designed this mosaic had observed
these animals not in their natural habitat, but in the arena of Pompeii’s amphitheatre. Animals were
sometimes forced to fight each other after having been starved and tormented to increase their
aggression. At other times, animals were part of “hunts” (venationes) or pitted against gladiators or
criminals condemned to die (damnatio ad bestias) [Author: MatthiasKabel. Distributed under a CC
BY-SA 2.0 license. Available online: https://cutt.ly/dRCLc99. Accessed: 2 February 2021]. (
b
) Plaster
cast of a pig found during the excavation in Villa Regina (Boscoreale). It certifies the breeding of pigs
on Vesuvian farms [Author: RealCarlo. Distributed under a CC BY 2.0 license. Available online:
https://cutt.ly/KRCLFFl. Accessed: 30 October 2021]. (
c
) Fresco of a rabbit and four figs (from
Herculaneum) [Author: Amphipolis. Distributed under a CC BY-SA 2.0 licence. Available online:
https://cutt.ly/3RCLM44. Accessed: 30 October 2021]. (
d
) Mosaic of marine life (from House of
the Geometric Mosaics,Pompeii) [Author: Carole Raddato. Distributed under a CC BY-SA 2.0 license.
Available online: https://cutt.ly/LRCZwkF. Accessed: 2 February 2021]. All images are of public
domain via Wikimedia Commons webpage.
Cattle were the most useful animals, producing milk, butter, cheese, meat, fertilizer,
horns and bones for tools, leather and fur for the manufacture of clothing [
120
]. Thus,
although cattle breeding was not mainly intended for meat consumption, owning a herd of
cattle was a sign of wealth for the Romans. During the winter, a good supply of cereals

Animals 2022,12, 213 10 of 37
was reserved for human consumption, for oxen and mules as pack animals to pull ploughs
in the field and for animals to transport and mill cereal grains [
114
,
117
,
120
]. Numerous
skeletal remains of cows, oxen and mules were found, mainly in the forum boarium in
Pompeii [
101
,
121
,
122
]. There are also numerous artistic representations in mosaics, frescoes
and sculptures showing cows, oxen and mules [117].
Horses were also used as a means of transportation, being mostly used for circus races
and army equites (knights) [
114
,
117
,
123
]. Their breeding was expensive and demanding, so
farmers preferred donkeys and mules, which were more resistant and needed less care [
123
].
Donkeys’ milk was highly appreciated, either for children—especially babies—or for use
as a base for cosmetics and therapeutic remedies. In addition, mowers and mills were
managed by these animals [
117
,
123
]. Various stables have been identified in Pompeii (e.g.,
House of Amarantus,House of Chaste Lovers, stables belonging to different bakeries) and
Herculaneum (e.g., a stable in Insula Orientalis II.1a belonging to a bakery on the north side of
Vicolo Meridionale), but their architecture and dimensions are generally not specific enough
to assign them, unambiguously, as rooms for oxen, horses or other livestock [
114
]. Five
equine skeletons were found in one of these stables, specifically in the House of Chaste Lovers
in Pompeii, another equid was found in Herculaneum [124], and recently three equids were
recovered in a large villa in Civita Giuliana, a suburb of Pompeii [
125
]. According to classical
taxonomic criteria and ancient mtDNA analysis, the classification of the Pompeian equine
remains found at the House of Chaste Lovers was identified as horses and
mules [126–129]
,
while the Herculanean remains were classified during the excavation phase and later
by genetic characterization, as those of a horse [
128
]. Once again, Roman art provides
representations of equines in different artistic works [117].
Sheep were the most widespread livestock because they provided meat for food, wool
for clothing, milk for cheese making and fertilizer for cultivating the fields. Goats were
also raised, as their milk was highly appreciated, and they also provided
leather [117,130].
Numerous bone and dental remains of sheep and goats were found in Pompeii and Hercula-
neum [101,121,122], and their representation in artistic works was frequent [117].
Pig farming was widespread, required little care and produced highly appreciated
meat [
114
,
122
]. Pliny the Elder (Naturalis Historia; 1st century CE) collected more than
50 recipes for pork, and similarly, several recipes were reported by Apicius in his gas-
tronomic treatise (De Re Coquinaria; 1st century CE). With their meat, they produced
long-lasting sausages, with the lard they cooked and with the suet they fed the oil lamps,
while with the bristles they made brushes. Numerous bone and dental remains of pigs were
found, mainly in the forum suarium of Pompeii [
101
,
121
,
122
], as well as other skeletal remains
uncovered during the excavations of the House of C. Julius Polybius in Pompeii [
131
]. Of
remarkable beauty is the plaster cast of a pig found in Villa Regina in Boscoreale (Figure 2b),
as well as different artistic works representing this animal [117,131].
Chickens and hens were also very widespread among farmed animals, these being
appreciated for their meat, as a supplier of eggs and their manure, which was an excellent
corrector of the most sterile vineyards and fields [
119
,
130
]; cocks, on the other hand,
were also bred as fighting animals and kept for sport [
117
]. According to Apicius (De
Re Coquinaria; 1st century CE), ducks and geese were also bred and highly appreciated,
especially for their eggs. In addition, bird feathers were used as ornaments for humans
and animals such as racehorses [
117
]. Interestingly, the carbonized remains of a chicken
were recovered on the top of an altar as a sacrificial offering in Pompeii [
46
]. As with all
other animals, Pompeii and Herculaneum provide innumerable artistic works depicting these
animals [117].
Rabbit and hare farming was appreciated for meat [
117
] (Figure 2c). According to
García [
132
] (p. 195), on display in the antiquarium of Pompeii prior to the 1943 WWII
bombing, was the skeleton of a rabbit that had been found near the oven of the Bakery and
shop of Sabinus.
In Roman times, beekeeping was widely practised to obtain wax and honey. The
Romans did not have sugar; although they produced a sweet syrup from beets, honey was

Animals 2022,12, 213 11 of 37
much more appreciated for its flavour, and it was even used for medical purposes [
133
].
Many ancient surviving Latin texts on agriculture and natural history provide important
information on beekeeping, such as those by Varro (Res Rusticae III.16.1–38; 1st century
BCE), Virgil (Georgics Book IV; 1st century BCE), Columella (De Re Rustica IX.2–16; 1st
century CE) and Pliny the Elder (Naturalis Historia XI.4–1; 1st century CE) [
134
]. Because
Roman hives were made from plant (biodegradable) materials, no material evidence is
known. However, five amphorae were discovered at Pompeii whose inscriptions were used
to label them (called tituli picti), indicating that they once held honey [133].
Aquaculture was a viable and highly productive industry in the Roman world, focused
on raising in piscinarii (artificial pools); either in freshwater or marine water, numerous
important aquaculture species were raised, such as moray eels, lobsters, sea breams, snap-
pers, sea bass, sturgeons, red mullets, meagres, barbels, pikes, octopus, eels, oysters and
mussels, among others [
117
,
135
]. According to Martial (Epigrammata X.30, 1st century CE),
the access to fresh, and even live, fish carried considerable social prestige [
136
], so with
time, incorporating a fishpond into the house or villa became an important status symbol.
More than 60 structures plausibly identified as fishponds or tanks have been discovered
in gardens and courtyards in Pompeii and Herculaneum [
137
]. Numerous remains of fish
and seafood from the Cardo V sewer were found in Herculaneum, showing a high diversity
of fish, marine mollusc and marine arthropod taxa [
64
]. There are also numerous artistic
works depicting many fresh and sea fish well attested as being farmed or fished by the
ancients, and frescoes and mosaics featuring scenes of seafood or of fishing [
137
]. Of
remarkable beauty are the mosaics from the House of the Faun, representing numerous types
of fish, and the mosaic of marine life from the House of the Geometric Mosaics, both in Pompeii
(Figure 2d).
The large mammals, already mentioned above, were raised mainly as a resource to
obtain food. In contrast, small mammals were considered unsuitable for human nutrition,
at least among Mediterranean peoples, who generally avoided feeding on rodents and
bats, as they were considered harmful to health [
138
]. However, the edible dormouse or
fat dormouse (Glis glis) represents an exception from this point of view (Figure 3a). In
the context of Roman gastronomy and, in particular, that of the imperial era, the edible
dormouse was considered a delicacy [
139
,
140
], as confirmed by Martial (Epigrammata III.58;
1st century CE), who narrates the unusual peasant tradition of offering edible dormouse
along with other products from the rural area as a sign of greeting and respect for the
neighbours [
138
,
141
], as well as different recipes transmitted by other Latin authors such
as Apicius (De Re Coquinaria VIII. 9, IX.1.1; 1st century CE) or Petronius (Satyricon XXXI.10;
1st century CE). During the II–I centuries BCE, a period in which the Roman world was
widely open to foreign influences, especially Greek and Eastern, refined ways of life were
overly promoted. This refinement of the society was fought in various ways by a sector
of the population that understood that its acceptance and dissemination threatened the
ideology and values, those of the traditional Roman, that had made the Roman Republic
great [
138
]. According to Pliny the Elder (Naturalis Historia VIII.223, XXXVI.4; 1st century
CE), in an attempt to curb the excessive pomp and idleness that prevailed in the social life
of patricians and manifested itself in luxury purchases and in the immoderate luxury of
banquets, parties and funerals, the most conservative senators promoted the enactment
of the so-called leges Sumptuariae (Sumptuary laws) that limited the expense that a citizen
could incur in various aspects of their day-to-day life. The lex Aemilia (78 BCE) stands out,
referring to the type and quantity of food that could be served at banquets. Among other
things, this law prohibited excessively refined cuisine, as well as the food use of oysters,
exotic birds and dormice [119,138,139].

Animals 2022,12, 213 12 of 37
Animals 2022, 12, x 12 of 38
the immoderate luxury of banquets, parties and funerals, the most conservative senators
promoted the enactment of the so-called leges Sumptuariae (Sumptuary laws) that limited
the expense that a citizen could incur in various aspects of their day-to-day life. The lex
Aemilia (78 BCE) stands out, referring to the type and quantity of food that could be served
at banquets. Among other things, this law prohibited excessively refined cuisine, as well
as the food use of oysters, exotic birds and dormice [119,138,139].
In addition to the historical and literary sources on the alimentary use of the
dormouse in Roman times, there are also numerous lines of evidence from archaeological
and zooarchaeological studies and research that confirm this practice [100,138–141]. Thus,
even though the dormouse was prohibited for consumption, it was still raised and used
as food by the Roman population. Skeletal and dental remains of dormouse have been
found in very small quantities, mainly at elite Roman sites, typically rural villages, but
also in suburban areas [100,101]. For example, dormouse bones were found in the forum
of Pompeii and in the north courtyard garden of the Villa of Poppaea in Oplontis [139,140].
However, their recovery, as with all microfauna remains, is highly dependent on effective
sieving campaigns during the bone retrieval phase. Although it is possible that the
dormice identified in these sites were eaten, there are no cut marks that corroborate this
directly [141]. The appreciation that the Romans had for the dormouse in the kitchen is
intimately linked to the peculiar breeding techniques in rural villages. As described by
Varro (Res Rusticae III.15.1–2; 1st century BCE), the dormice were reared and fattened in
earthenware containers [140], called vivaria in doliis by Pliny the Elder (Naturalis Historia
VIII.211,224; 1st century CE) [141] (Figure 3b). The internal surface of these dolia was
characterized by the presence of parallel and concentric protrusions—called semitae—or
by a single continuous helical protrusion through which the animals could move without
getting dirty with their own faeces that accumulated at the bottom of the container. The
surface of these vivaria in doliis (also known as gliraria) had many small holes, presumably
to aerate the container. In addition, the openings connected to small containers would
probably allow the animals to be provided with drinking water and/or food [119,140].
Several gliraria have b een foun d, but on ly nine c an, wi th reaso nable ce rtainty, be iden tified
as such. Although some of them were found in different Roman provinces, six were found
around the area of Mt. Somma–Vesuvius: four in Pompeii, one in Herculaneum and one in
Boscoreale. The fact that gliraria were mainly found in villae suggests that the consumers of
dormouse were predominately members of the Roman elite [140].
Figure 3. (a) Edible dormouse (Glis glis), considered a delicacy by the Romans [Author: Azay.
Distributed under a CC BY-SA 2.5 license. Available online: https://cutt.ly/bRCLi6u. Accessed: 30
October 2021]. (b) Reconstruction of a vivarium in doliis (also known as glirarium) at the Museum of
Figure 3.
(
a
) Edible dormouse (Glis glis), considered a delicacy by the Romans [Author: Azay.
Distributed under a CC BY-SA 2.5 license. Available online: https://cutt.ly/bRCLi6u. Accessed:
30 October 2021]. (
b
) Reconstruction of a vivarium in doliis (also known as glirarium) at the Museum
of the Etruscan Academy in Cortona. The vivarium in doliis is a terracotta container used for keeping
edible dormouse. By inducing hibernation via darkness and confinement, the vivarium in doliis would
cause the dormouse to fatten [Author: Viscondedeportoseguro. Distributed under a CC BY-SA 4.0
license. Available online: https://cutt.ly/BRCZdek. Accessed: 30 October 2021]. All images are of
public domain via Wikimedia Commons webpage.
In addition to the historical and literary sources on the alimentary use of the dor-
mouse in Roman times, there are also numerous lines of evidence from archaeological
and zooarchaeological studies and research that confirm this practice [
100
,
138
–
141
]. Thus,
even though the dormouse was prohibited for consumption, it was still raised and used
as food by the Roman population. Skeletal and dental remains of dormouse have been
found in very small quantities, mainly at elite Roman sites, typically rural villages, but
also in suburban areas [
100
,
101
]. For example, dormouse bones were found in the forum
of Pompeii and in the north courtyard garden of the Villa of Poppaea in Oplontis [
139
,
140
].
However, their recovery, as with all microfauna remains, is highly dependent on effec-
tive sieving campaigns during the bone retrieval phase. Although it is possible that the
dormice identified in these sites were eaten, there are no cut marks that corroborate this
directly [
141
]. The appreciation that the Romans had for the dormouse in the kitchen is
intimately linked to the peculiar breeding techniques in rural villages. As described by
Varro (Res Rusticae III.15.1–2; 1st century BCE), the dormice were reared and fattened in
earthenware containers [
140
], called vivaria in doliis by Pliny the Elder (Naturalis Historia
VIII.211,224; 1st century CE) [
141
] (Figure 3b). The internal surface of these dolia was
characterized by the presence of parallel and concentric protrusions—called semitae—or
by a single continuous helical protrusion through which the animals could move without
getting dirty with their own faeces that accumulated at the bottom of the container. The
surface of these vivaria in doliis (also known as gliraria) had many small holes, presumably
to aerate the container. In addition, the openings connected to small containers would
probably allow the animals to be provided with drinking water and/or food [
119
,
140
].
Several gliraria have been found, but only nine can, with reasonable certainty, be identified
as such. Although some of them were found in different Roman provinces, six were found
around the area of Mt. Somma–Vesuvius: four in Pompeii, one in Herculaneum and one in
Boscoreale. The fact that gliraria were mainly found in villae suggests that the consumers of
dormouse were predominately members of the Roman elite [140].

Animals 2022,12, 213 13 of 37
4.3.3. Household Animals
The Romans benefited from collaboration and companionship with different animals,
and both then and now, the dog was their favourite companion animal [
142
]. According to
Virgil (Georgics III.404–413; 1st century CE), Columella (De Re Rustica VII.11.1–2; 1st century
CE), Strabo (Geographica XV.1; 1st century CE), Grattius (Cynegetica 179; 1st century CE)
and Pliny the Elder (Naturalis Historia VIII.61; 1st century CE), in ancient Rome, the dog
performed various functions [
117
,
143
–
145
]. On the one hand, it was used in popular shows
and in the largest circuses, where they fought against each other or against aggressive bears,
tigers and lions [
117
]. For this, the molossus was used, a canid with a strong complexion
endowed with piercing fangs and a strong jaw, as described by Virgil (Georgics III.404–413;
1st century CE) and Varro (Rerum Rusticarum II.9; 1st century BCE). Seeing its potential in
shows, the molossus was also introduced in warlike contexts, constituting the so-called canes
pugnaces (fighting dogs). They were brought to the battlefield in packs of several dozen by
special units of the Roman legion [143].
On the other hand, dogs were also highly prized on hunts. The Romans called hunting
dogs canes venatici (sporting dogs) and divided them according to the kind of hunting for
which they were predestined: the sagaces were those used to follow the tracks of the prey
(hounds); the celeres were the fast dogs used for the pursuit of the prey, with a special
preference for the greyhound (canes vertragus); and the already commented canes pugnaces,
used to attack prey, especially wild boars and other wild animals. The rural environment
was another place where dogs stood out, also being used for herding or guarding the
villages. The canes pastorales (shepherd dogs) were used for the care and transport of
livestock. In addition, they defended livestock from predators that lurked on the roads
and in the fields and forests. The canes villatici (watchdogs) were the dogs destined for the
custody of domus,insulae,villae or workshops, warning if strangers appeared [143,146].
At the end of the Republican period (during the 1st century BCE), the keeping of dogs
to serve as guardians of the home became fashionable among the wealthier Roman classes.
Thus, in some Roman domus, we find represented in mosaics how these ‘companion’ dogs
served as a warning to intruders. They were represented with the inscription ‘Cave canem’
(Beware of the dog!). As an example, we can highlight the mosaic at the entrance door to
the House of the Tragic Poet (Figure 4a) or the mosaic in the foyer of the House of Paquius
Proculus, both in Pompeii [
117
]. It was common for these mosaics to represent a molossus, a
dog whose impressive musculature impressed and intimidated anyone entering the domus.
The plebeians, generally with scarce economic resources, could not afford this type of
guardian animal, so they had geese instead. Their territorial character, along with their
loud squawk, made them an excellent and inexpensive guardian.
In a society as refined as the Roman one, the dog also became a much-loved companion.
To serve as pets, it was enough to be docile and faithful, like the small canis catelli. Their
sole purpose was to be an entertainment for children, a flea and fly deterrent for its owners
and a symbol of social status for wealthy women. This class of dogs (lapdogs) were so
appreciated that the description that the Roman poet Martial (Epigrammata I.109; 1st century
CE) makes of the dog named Issa of his friend Publio reflects it clearly [147]. Martial says:
“[
. . .
] Issa is more pure than the kiss of a dove. Issa is more loving than any maiden. Issa is
dearer than Indian gems. [
. . .
] That her last hour may not carry her off wholly, Publius has her
limned in a picture, in which you will see an Issa so like, that not even herself is so like herself.”
(translated by [
148
]). However, these pet dogs were not common among the plebeians
because, generally, having a dog that did not work—either as a guardian, shepherd or
hunter—was beyond most people’s comprehension.
Numerous ancient Latin texts and artistic representations in mosaics, frescoes bas-
reliefs, statues and other archaeological artefacts evidence the presence and diffusion of
those types of dogs [
117
]; for example, the mosaics depicting watchdogs found in the
Houses of Paquius Proculus,Vesonius Primus and the House of the Tragic Poet in Pompeii; the
mosaics (House of the Wild Boar), frescoes (House of the Gladiators,House of the Wild Boar,
House of Menander) and statues (Domus of L. Rapinasi Optati, House of the Deers) depicting

Animals 2022,12, 213 14 of 37
hunting scenes in Pompeii and Herculaneum; and archaeological remains of what are consid-
ered doghouses in the House of Achilles and the House of the Garden of Hercules in Pompeii.
Furthermore, numerous skeletal remains of dogs have been found in Pompeii, such as, for
example, those found in the House of Amarantus,House of Paquius Proculus,House of Romulus
and Remus and Villa of Diomedes in Pompeii. Interestingly a plaster cast of a watchdog was
found in the House of Vesonius Primus in Pompeii, showing his leather collar with the two
bronze rings for the chain (Figure 4b).
Animals 2022, 12, x 14 of 38
among the plebeians because, generally, having a dog that did not work—either as a
guardian, shepherd or hunter—was beyond most people’s comprehension.
Figure 4. (a) Mosaic at the entrance door to the House of the Tragic Poet (Pompeii) depicting a dog
served as a warning to intruders, with the inscription ‘Cave canem’ (Beware of the dog!) [Author:
Sailko. Distributed under a CC BY-SA 4.0 license. Available online: https://cutt.ly/JRCZnid.
Accessed: 30 October 2021]. (b) Plaster cast of a watchdog found in the House of Vesonius Primus
(Pompeii), showing his leather collar with the two bronze rings for the chain [Author: Jebulon.
Distributed under a CC0 1.0 license. Available online: https://cutt.ly/TRCZmFG. Accessed: 30
January 2021]. (c) Mosaic from the House of the Faun (Pompeii) representing a cat attacking a partridge
[Author: Marie-Lan Nguyen. Distributed under a CC-PD-Mark 1.0 license. Available online:
https://cutt.ly/ARCZQYm. Accessed: 30 January 2021]. (d) Mosaic from the House of the Faun
(Pompeii) depicting parrots, a dove and a cat [Author: Marie-Lan Nguyen. Distributed under a CC-
PD-Mark 1.0 license. Available online: https://cutt.ly/wRCZWLt. Accessed: 30 January 2021]. All
images are of public domain via Wikimedia Commons webpage.
Numerous ancient Latin texts and artistic representations in mosaics, frescoes bas-
reliefs, statues and other archaeological artefacts evidence the presence and diffusion of
those types of dogs [117]; for example, the mosaics depicting watchdogs found in the
Houses of Paquius Proculus, Vesonius Primus and the House of the Tragic Poet in Pompeii; the
mosaics (House of the Wild Boar), frescoes (House of the Gladiators, House of the Wild Boar,
House of Menander) and statues (Domus of L. Rapinasi Optati, House of the Deers) depicting
hunting scenes in Pompeii and Herculaneum; and archaeological remains of what are
considered doghouses in the House of Achilles and the House of the Garden of Hercules in
Pompeii. Furthermore, numerous skeletal remains of dogs have been found in Pompeii,
such as, for example, those found in the House of Amarantus, House of Paquius Proculus,
House of Romulus and Remus and Villa of Diomedes in Pompeii. Interestingly a plaster cast of
a watchdog was found in the House of Vesonius Primus in Pompeii, showing his leather
collar with the two bronze rings for the chain (Figure 4b).
The cat is another animal found in the Roman domestic environment. There is some
evidence of coexistence between cats and men as early as ancient Egypt—where they were
traditionally venerated as a sacred animal—and later also in Greece, although there this
Figure 4.
(
a
) Mosaic at the entrance door to the House of the Tragic Poet (Pompeii) depicting a dog served
as a warning to intruders, with the inscription ‘Cave canem’ (Beware of the dog!) [Author: Sailko.
Distributed under a CC BY-SA 4.0 license. Available online: https://cutt.ly/JRCZnid. Accessed:
30 October 2021]. (
b
) Plaster cast of a watchdog found in the House of Vesonius Primus (Pompeii),
showing his leather collar with the two bronze rings for the chain [Author: Jebulon. Distributed
under a CC0 1.0 license. Available online: https://cutt.ly/TRCZmFG. Accessed: 30 January 2021].
(
c
) Mosaic from the House of the Faun (Pompeii) representing a cat attacking a partridge [Author:
Marie-Lan Nguyen. Distributed under a CC-PD-Mark 1.0 license. Available online: https://cutt.ly/
ARCZQYm. Accessed: 30 January 2021]. (
d
) Mosaic from the House of the Faun (Pompeii) depicting
parrots, a dove and a cat [Author: Marie-Lan Nguyen. Distributed under a CC-PD-Mark 1.0 license.
Available online: https://cutt.ly/wRCZWLt. Accessed: 30 January 2021]. All images are of public
domain via Wikimedia Commons webpage.
The cat is another animal found in the Roman domestic environment. There is some
evidence of coexistence between cats and men as early as ancient Egypt—where they were
traditionally venerated as a sacred animal—and later also in Greece, although there this
animal never gained the prestige it had in Egypt [
114
,
149
]. It was in the 1st century CE
when the cat would begin to be known in the Roman world. The Romans used colubers and
mustelids, such as weasels, stone martens and ferrets, for the control of rodents [
142
,
149
];
but they soon realized that cats were more easily tamed and appreciated the services
both as a working animal—to eradicate rodents from their homes—and as a companion
animal. Therefore, according to Pliny the Elder (Naturalis Historia X.73; 1st century CE),
domestic cats replaced mustelids at the beginning of the Christian era [
150
]. During the

Animals 2022,12, 213 15 of 37
conquest campaigns, the Romans took cats with them, contributing to its spread throughout
Europe [
149
]. The reason they took them was so that they would hunt the many rodents in
the camps and winter quarters. In addition, after passing through Egypt, due to its link
with Isis, the cat was considered an animal that symbolized victory. Traces of the presence
of the cat have been found in all the regions conquered by the Romans [105,149,150].
Although the skeletal remains of many diverse animals abounded in Pompeii and
Herculaneum, the skeletal remains of cats are extremely rare. The skeletal remains of a cat
have only been found in a large commercial vineyard at Pompeii [
151
] and in the forum
(cited in [
101
]). However, in Pompeii, there is no shortage of representations of the feline: of
remarkable beauty, for example, is the mosaic from the House of the Faun that depicts a cat
attacking a partridge, now kept in the Naples National Archaeological Museum (Figure 4c).
The same museum exhibits another Pompeian mosaic depicting parrots, a dove and a cat,
also from the House of the Faun [
117
] (Figure 4d). In any case, it should not be forgotten
that Pompeii had many contacts with Greece and Egypt; that is why it is believed that the
‘Pompeian’ cats, the few that there were, managed to save themselves at the first signs of
the catastrophe [117]. However, these are just mere speculation.
4.3.4. Wild Animals
The presence of loose animals in the city was a constant and was part of the ancient
Roman urban landscape. Dogs were abundant even in the homes of the poorest people, not
only as guardians of modest homes, but also used to dispose of food scraps (as pigs had
also been used) [
114
]. Tanners, for their part, hunted them to make leather, and the most
disadvantaged people probably consumed meat from stray dogs as a source of protein in
the absence of other prey, this being an atypical way of acquiring hydatid cysts [85].
However, there were many other animals that roamed the public roads of Roman
cities. Horace (Epistulae II.2. 72–75; 1st century BCE) describes the city of Rome as noisy and
chaotic, emphasizing how busy the streets were and the lack of order when people and ani-
mals made their way down the street together. As an example, Horace mentions a muddy
sow walking calmly through the city [
152
]. At that time, there were no slaughterhouses as
we know them today, but the animals were bought in the corresponding livestock market
(forum boarium: cattle market; forum suarium: pig market) and driven alive, urinating and
defecating in the streets, to the macellum (public market dedicated to the sale of meat and
fish market) and specialized butchers’ shops, where they were slaughtered, disembowelled,
dismembered and sold. Undoubtedly, the continuous traffic of animals such as pigs, sheep
and cattle contributed to the general dirtying of the streets with excrement. In addition,
leftovers were often dumped directly onto the street, which, together with the low levels of
sanitation and poor standards of public health, implied a high risk of numerous diseases
such as rabies, bovine tuberculosis, parasitosis (e.g., scabies, echinococcosis, trichinellosis,
taeniasis), ringworm, brucellosis, leptospirosis and many more. If the frequent faecal con-
tamination of water and food is also included, this list would be extended to other diseases,
such as cholera, hepatitis, dysentery, typhoid, salmonellosis, colibacillosis, etc. [85,114].
As for rodents, the black rat (Rattus rattus) would be the protagonist, already men-
tioned above (see Section 4.2. Trade routes). However, we can also highlight the wood
mouse (Apodemus sylvaticus). The wood mouse, like the house mouse (Mus musculus) (see
Section 4.1. Population density and housing), is an agricultural pest that damages cereals,
orchards, pastures, horticultures and tubers [
95
,
153
]. However, unlike the house mouse,
the wood mouse can survive in environments where humans are not present, allowing it to
colonize a wider range of environments and travel between human and wild environments.
Thus, this higher level of mobility could have influenced a greater spread of diseases.
The spread of diseases transmitted by rodents is not only influenced by available food
and environmental conditions, including pathogens and disease vectors present in the
environment, but also by factors specific to the species itself, such as the population density,
the size of the distribution area, and the variety of environments that the species is capable
of inhabiting [
154
]. Thus, wood mice are generally more dangerous vectors of diseases

Animals 2022,12, 213 16 of 37
than house mice. Their greater range and greater flexibility to the habitat in which they live
provide them with greater opportunities to contract diseases or contract disease-bearing
parasites and a greater ability to transmit these diseases and parasites over long distances.
A contemporary example would be the transmission of the pulmonary syndrome by the
hantavirus, a dangerous respiratory infection that is transmitted indoors when wild rodents
invade them in bad weather situations [155].
House mice frequently outnumber wild mice, suggesting that a high density of house
mice in an ancient city may have had the additional effect of protecting the human pop-
ulation from the importation of external diseases by wild mice [
100
]. If Pompeii and
Herculaneum had not been occupied by a high density of house mice (Mus musculus), as
shown by the amount of skeletal remains found in these cities, they would probably have
been occupied by wood mice (Apodemus sylvaticus), as suggested by the evidence of the
different excavations carried out [100].
Summarizing, animals—whether livestock, wild, exotic or household animals—
accompanied the Romans in most of their daily chores, constituting a health risk for people
in close contact with them. Thus, bites, scratches and allergies would be the most common
threats and would result in localized infections; however, there are other infections—
parasitic, bacterial, viral and fungal—that could be transmitted by direct contact with these
animals, their urine and excrement or through arthropods, such as dogs (e.g., rabies, hydati-
dosis, toxocariosis, leptospirosis, asthma), cats (e.g., toxoplasmosis, toxocariosis, cat scratch,
rabies, pasteurella), rodents (e.g., hantavirus, leptospirosis, salmonellosis, leishmaniasis,
toxoplasmosis), pigs (e.g., cysticercosis, taeniasis, balantidiasis) or birds (e.g., psittacosis,
cryptococcosis, histoplasmosis, avian flu), among others (e.g., [98,99,114,156,157]).
4.4. Occupational and Work-Related Zoonotic Diseases
After the Social War (91–87 BCE), Pompeii was elevated to the status of Roman colony
under the name of Colonia Cornelia Veneria Pompeianorum (80 BCE), whereas Herculaneum
received the lower rank of municipium (89 BCE) [
158
]. This situation led to an enormous
development of both cities, both from an urban and economic point of view. There was a
great commercial boost thanks to the fertility of the volcanic soil and its excellent geograph-
ical position in the centre of the Gulf of Naples. Thus, commerce played a fundamental
role in the local economy, as evidenced by the abundance of shops and taverns unearthed
in Pompeii and Herculaneum. Mainly agricultural products were commercialized, although
artisanal or “industrial” activities related to the transformation of farm products were also
developed [43,44].
The historical sources, particularly from the I–II centuries CE, bear witness to the great
variety of commercial activities, occupations and trades in these ancient Roman cities [
44
].
Economic activities of all kinds were located throughout the city, especially along the main
arteries of both cities: the Via dell’Abbondanza in Pompeii and the Decumanus Maximus in
Herculaneum. Among all the commercial activities, some of them represented certain risks,
with some specific activities and trades that increased the probability of zoonotic infection.
These zoonotic occupational diseases are common when there is close contact at work with
animals or animal products. The main occupations with a risk of zoonosis were those in
which workers were in contact with: (i) infected animals; (ii) materials or products from
infected animals; (iii) secretions from infected animals, including saliva, blood and faeces;
(iv) aerosols or powders contaminated with secretions from infected animals; (v) soil or
water contaminated with materials or secretions from infected animals; or (vi) infected
vectors implicated in the transmission of a zoonosis [
159
,
160
]. The occupations with the
highest risk of zoonotic infections were those of the agricultural industry, as farmers had
close contact with potentially infected animals during land cultivation activities. Other
hazardous occupations with close contact with animals included those involved in animal
husbandry (e.g., aviaries), those involved in the animal trade (e.g., importing exotic animals
for public games), activities of handling and processing of potentially infected animal
products, as well as those activities related to the cleaning and hygiene of the facilities

Animals 2022,12, 213 17 of 37
where animals or animal products were kept. Certain zoonoses are particularly associated
with contaminated water, so the risk of infection was high in occupations that were in close
contact with water, including certain occupations related to wastewater treatment.
4.4.1. Agriculture
Agriculture was the basis of the Roman economy, whose main concern was feeding the
vast number of citizens and legionaries who populated the Mediterranean
region [44,161]
.
However, with regard to excess human waste, growing cities such as Pompeii and Her-
culaneum began to have problems. This situation encouraged the Romans to find other
ways not only to dispose of this progressively accumulating waste, but also to use them for
commercial exploitation [162].
Human urine was not only used for mordanting certain dyestuffs and for cleaning
purposes [
85
,
163
] (see Section 4.4.3. Textile industry: Laundries (fullonicae) for more details),
but also, because it contains significant levels of nitrogen, phosphorus and potassium, it
met the requirements for an efficient fertilizer for crop cultivation [
164
]. For example, the
ancient Roman writer expert in agriculture Columella (De Re Rustica II.15, 1st century CE)
asserts that “[The] human urine, which you have let grow old for six months, is fitter for shoots
and young trees. If you apply it to vines, or to apple-bearing trees, there is nothing that contributes
more to make them bear abundance of fruit; nor does this thing only produce a greater increase, but
it also improves both the taste and the flavour of the wine, and of the apples.” (translated by [
165
]).
Columella (De Re Rustica II.23, 1st century CE) also suggests watering pomegranate trees
with human urine, making them juicier and tastier [166].
Another commercial exploitation involving human waste was the use of excrement
as fertilizer. In this regard, in the Roman world, human excrement was used as a comple-
mentary or substitute of animal manure to fertilize the soil of croplands [
162
]. Columella
(De Re Rustica I.6.24, 1st century CE) promoted the use of mixed animal and human ex-
crement (stercorata) since he noticed that manure from both barnyard and thermae (baths)
worked best [
167
]. The Romans believed that nature did not admit waste and included
everything natural in a spiral of life and death. In this way, it can be said that the Romans
had realized that recycling the bodily waste of humans and animals was a good thing for
the productivity of the soil [
167
]. Cato the Elder (De Agri Cultura LXI, 2nd century BCE)
confirms the importance of manure to nourish croplands: “What is the first principle of good
agriculture? To plough well. What is the second? To plough again; and the third is to manure
[. . . ]
” (translated by [
168
]). Other Latin author experts in agriculture, such as Varro and
Columella, attest to the practice of using human excrement to manure the croplands. Varro
(Res Rusticae I.38.1–3; 1st century BCE) informs that “[
. . .
] several kinds [of manure] have
different qualities.”, and states that “[
. . .
] the best manure is that of birds, but the best of all is
[
. . .
] the manure of pigeons because it is the hottest and causes the land to ferment. [
. . .
] The
manure next in value to that of doves is human faeces, and third that of goats and sheep and asses.
The manure of horses is of the least value on corn land [
. . .
].” (translated by [
168
]). On his part,
Columella (De Re Rustica II.15; 1st century CE) asserts that “There are three principal sorts of
dung: that produced by fowls, that by men, and that by cattle. Of fowls that [which] is reckoned
the best, which is brought out of pigeon-houses. [
. . .
] The second is that which men make, if it be
mixed with the other filth and sweepings of the manor-house; for it is naturally very hot by itself,
and therefore burns up the ground.” (translated by [165]).
The literary sources verify that the Romans used animal and human waste in agricul-
ture, but apparently, human waste was not the first choice, despite its copious availability
in urban centres (see Section 4.5. Latrines, sewers and baths (thermae) for more details),
but it was frequently used as a supplement to animal manures [
85
,
167
]. It is easy to under-
stand how the use and management of human faeces as fertilizer could have constituted a
danger to public health as these, not being subjected to any specific disinfectant treatment,
could have kept various pathogens within them which, once dissolved, would infiltrate
the soil and aquifers and then consequently contaminate the foods through which it was
easy for the population to become reinfected [
169
,
170
]. Edible vegetables and fruits fertil-

Animals 2022,12, 213 18 of 37
ized with human waste held considerable risks and could be contaminated with various
viruses, bacteria and worm eggs. Moreover, if disease carriers such as insects, rodents
and pets came into contact with excrement, especially human excrement, and then came
near food or drinking water, the problem could become chronic [
85
,
167
]. Scobie [
85
] dis-
cusses the various diseases to which Romans were susceptible in the light of their usage
of human excrement as fertilizer (e.g., cholera, dysentery, gastroenteritis, infectious hep-
atitis, leptospirosis, typhoid fever) and lists innumerable common genera of pathogenic
organisms that can be found in water contaminated by infected faeces (e.g., salmonellosis,
colibacillosis).
4.4.2. Livestock Farming
As already mentioned above (see Section 4.3.2. Livestock), livestock farming and pas-
toralism were among the main economic resources of Roman cities, being related to a series
of activities in which livestock were handled. Within these activities, the operations that
would produce a transmission of zoonotic diseases would be, for example, (i) assistance to
the birthing of animals; (ii) care of the offspring; (iii) feeding of livestock; (iv) transportation
of livestock; (v) milking of dairy breeds (i.e., goats, sheep, cows); (vi) sheep shearing;
(vii) hooves’ care; (viii) amputation of bovine antlers; (ix) caring for sick animals; and
(x) cleaning of animals and their facilities [
171
]. Often, during these activities, Roman
workers had to stay with the animals or even grab them, which would represent a series of
risks that would materialize in the form of accidents whose origin would be found in the
unpredictable reaction that the animals could have (e.g., scratches, bites).
4.4.3. Textile Industry
After agriculture, one of the most important artisanal economic activities was the cloth-
ing industry, a large, highly profitable and highly organized trade in different sectors [
172
].
Tanneries (officina coriariorum). The tanning industry was the industrial sector that
produced hides and skins by recovering and enhancing a by-product of the food industry:
raw animal skin from slaughter. The officina coriariorum of M. Vesonius Primus is the only
tanning complex identified in Pompeii, where the rawhides were macerated and processed
to be transformed into leather. The tannery was operational in 79 CE and was divided
into separate compartments necessary for all stages of the tanning process [
173
]: in the
courtyard, there was a stone bench for skinning the animals, and the courtyard was also
used for the storage of fresh leathers to be treated; the tanning of fresh leathers was carried
out in a complex of tanks arranged under a portico; under the arcades of the courtyard
there were the systems for tanning leather, finishing, cutting and making leather objects.
Furthermore, in this tanning complex were found tools characteristic of leather processing,
suitable for each of the stages of tanning: a semi-circular blade knife used to epilate and
deflesh the hides, a tool with a rectangular metal blade to soften and smooth the leather, and
a circular blade knife used by tanners and shoemakers to pre-cut pieces of leather [
173
,
174
].
In ancient Pompeii, the tanning industry was closely linked to that of wool; in fact, M.
Vesonius Primus was not only the manager of a tannery but also of a fullonica (industry
for cleaning clothes) among the many that were present in the rich Vesuvian town (see
Section 4.4.3. Textile industry: Laundries (fullonicae) for more details).
Tannery workers were at considerable risk of developing occupational contact der-
matitis, and this inflammation of the skin could constitute the main danger to health caused
by contact of the skin and certain chemicals and tools used in the handling and elaboration
of raw hides and skins [
175
]. Other health risks could be carbuncle or anthrax, an infection
caused by the bacterium Bacillus anthracis, the main risk being contact with skin or hair,
bone products and wool or with infected animals. Inhalation of anthrax could occur if
anthrax spores entered the lungs through the respiratory tract. This type of infection can be
contracted when workers inhale airborne anthrax spores during processes such as leather
tanning and wool processing [40,176].

Animals 2022,12, 213 19 of 37
Laundries (fullonicae).
Theoretically, five types of textile workshops can be distin-
guished: (i) initially raw wool was washed and combed in the officinae lanifricariae; (ii) then
the wool was dyed in the officinae tinctoriae; (iii) spinning and weaving were carried out
in the textrinae; (iv) the wool was felted in the officinae coactiliariae; and (v) the fullonicae
were the establishments where the finishing of woollen clothes took place, but which also
took on the functions of a laundry service [
177
]. However, this distinction, based on the
archaeological record, often does not allow a clear differentiation between the different
types and all of them are named with the general term fullonicae [178] (Figure 5a).
Animals 2022, 12, x 19 of 38
of leather [173,174]. In ancient Pompeii, the tanning industry was closely linked to that of
wool; in fact, M. Vesonius Primus was not only the manager of a tannery but also of a
fullonica (industry for cleaning clothes) among the many that were present in the rich
Vesuvian town (see Section 4.4.3. Textile industry: Laundries (fullonicae) for more details).
Tannery workers were at considerable risk of developing occupational contact
dermatitis, and this inflammation of the skin could constitute the main danger to health
caused by contact of the skin and certain chemicals and tools used in the handling and
elaboration of raw hides and skins [175]. Other health risks could be carbuncle or anthrax,
an infection caused by the bacterium Bacillus anthracis, the main risk being contact with
skin or hair, bone products and wool or with infected animals. Inhalation of anthrax could
occur if anthrax spores entered the lungs through the respiratory tract. This type of
infection can be contracted when workers inhale airborne anthrax spores during processes
such as leather tanning and wool processing [40,176].
Laundries (fullonicae). Theoretically, five types of textile workshops can be
distinguished: (i) initially raw wool was washed and combed in the officinae lanifricariae;
(ii) then the wool was dyed in the officinae tinctoriae; (iii) spinning and weaving were
carried out in the textrinae; (iv) the wool was felted in the officinae coactiliariae; and (v) the
fullonicae were the establishments where the finishing of woollen clothes took place, but
which also took on the functions of a laundry service [177]. However, this distinction,
based on the archaeological record, often does not allow a clear differentiation between
the different types and all of them are named with the general term fullonicae [178] (Figure
5a).
Figure 5. (a) Fullonica of Stephanus (Pompeii), showing the atrium with laundry basin [Author:
WolfgangRieger. Distributed under a CC BY-SA 4.0 license. Available online:
https://cutt.ly/xRCZOId. Accessed: 30 October 2021]. (b) Roman fresco from the Fullonica of Lucius
Veranius Hypsaeus (Pompeii). The man on the left is busy brushing wool cloth. The man on the right,
standing beneath a caged dome, is engaged in fabric whitening via sulphurized fumigation. An owl
is perched on top of the cage, likely a symbol of Athena, protector of the lanaiuoli (i.e., companies of
wool merchants) [Author: WolfgangRieger. Distributed under a CC-PD-Mark 1.0 license. Available
online: https://cutt.ly/DRCZPWP. Accessed: 30 October 2021]. All images are of public domain via
Wikimedia Commons webpage.
The Fullonica of Lucius Veranius Hypsaeus in Pompeii exemplifies the arrangement and
furnishing of a Pompeian fullones’ establishment, in which depictions of a textile press and
work scenes were found in the inner courtyard (Figure 5b). These iconographic
representations give us important information about the production sequence. The first
operation was that of washing, which was done with “fuller’s earth”—a type of clay
mixed with natural alkaline chemicals such as carbonate of soda, potash and human
urine—to remove any oily material. This was done in tubs filled with a cleaning solution
Figure 5.
(
a
)Fullonica of Stephanus (Pompeii), showing the atrium with laundry basin [Author:
WolfgangRieger. Distributed under a CC BY-SA 4.0 license. Available online: https://cutt.ly/
xRCZOId. Accessed: 30 October 2021]. (
b
) Roman fresco from the Fullonica of Lucius Veranius
Hypsaeus (Pompeii). The man on the left is busy brushing wool cloth. The man on the right, standing
beneath a caged dome, is engaged in fabric whitening via sulphurized fumigation. An owl is perched
on top of the cage, likely a symbol of Athena, protector of the lanaiuoli (i.e., companies of wool
merchants) [Author: WolfgangRieger. Distributed under a CC-PD-Mark 1.0 license. Available online:
https://cutt.ly/DRCZPWP. Accessed: 30 October 2021]. All images are of public domain via
Wikimedia Commons webpage.
The Fullonica of Lucius Veranius Hypsaeus in Pompeii exemplifies the arrangement and
furnishing of a Pompeian fullones’ establishment, in which depictions of a textile press
and work scenes were found in the inner courtyard (Figure 5b). These iconographic
representations give us important information about the production sequence. The first
operation was that of washing, which was done with “fuller’s earth”—a type of clay mixed
with natural alkaline chemicals such as carbonate of soda, potash and human urine—to
remove any oily material. This was done in tubs filled with a cleaning solution and cleaned
by the fullones (laundry workers) treading upon them. When dry, the cloth was brushed
and carded to raise the nap to make the surface even. The cloth was then fumigated with
sulphur and bleached in the sun while spread out over a frame. Cleaning soiled clothes was
another function of the fullones [
58
,
172
]. Science tells us that bacteria transform the urea
present in urine into ammonia [
179
]. The Roman poet Catullus (Carmina 39.20, 1st century
BCE) could not have known this, but he certainly knew the whitening power of urine on
the teeth, of which he expressly speaks in one of his Carmina, a sign that it was a practice
adopted by his contemporaries [
180
]. However, ammonia has degreasing properties and
fixes natural colours, so human urine was also used as a detergent, as a mordant for some
dyes and as a bleach in the textile industry [173].
The collection and use of urine by fullones for mordanting certain dyestuffs reveal an
area of private enterprise in the disposal and commercial exploitation of human waste in
Roman cities [
85
]. Thirty-nine fullonicae were active in Pompeii, for which large quantities of
urine were indispensable. That urine was widely used at that time is demonstrated by the

Animals 2022,12, 213 20 of 37
story of the famous vectigal urinae (the urine tax), which tanners and fullones were forced
to pay on the urine collected in public latrines. It was imposed by Emperor Vespasian,
aware of the vast use of urine in the textile industry. The Roman historian Suetonius (De
vita Caesarum, Vespasian 23.3, 2nd century CE) reports that when Vespasian’s son Titus
complained about the disgusting nature of the tax, his father held up a gold coin and asked
whether he felt offended by its smell (sciscitans num odore offenderetur). When Titus said
“No”, Vespasian replied, “Yet it comes from urine” (Atqui ex lotio est) [179].
As can be imagined, being a fullo was certainly not a good job, and even living near
these commercial buildings was not supposed to be pleasant. To get the urine needed, the
fullonicae encouraged pedestrians to supply urine through jars hung on the wall placed in
front of these shops [
162
,
181
]. References to the olfactory nuisance caused by fulling are
few and fragmentary. Martial (Epigrammata VI.93.1–2) associates fullonicae with bad smells
in an epigram, whose text invokes a situation in which one of the overused dolia curta
(urine container) (see Section 4.5. Latrines, sewers and baths (thermae) for more details) was
accidentally broken, either knocked over in the street or during transport by the fullones
and the odour circulated freely through the air [
182
]. In any case, since the cloths at the
end of the procedure were clean and odourless, the practice of recycling urine and its
industrial use continued for centuries until chemistry was able to synthesize urea and
produce synthetic detergents. Detailed information on the health implications because
of the handling of human faeces and urine is shown in Section 4.5. Latrines, sewers and
baths (thermae).
4.4.4. Storage and Grain Processing, and Bakeries (Pistrina)
Ancient literary sources (e.g., Cato de Elder De Agri Cultura, 2nd century BCE; Varro
Res Rusticae Book III, 1st century BCE; Columella De Re Rustica, 1st century CE; Pliny the
Elder Naturalis Historia, 1st century CE; Apicius De Re Coquinaria, 1st century CE), as well
as archaeological and biological evidence, show that legumes (e.g., faba bean, lablab bean,
cowpea, lentil, chick pea, common pea, grass pea, white lupine, bitter vetch, fenugreek,
mung bean, alfalfa), together with cereals (e.g., wheat, barley, sorghum, durum, einkorn,
emmer, millet, oats, rice, rye), constituted most of the nutritional and energy needs of an
individual in ancient times [
183
–
186
]. As the Roman Empire expanded, huge amounts of
food were needed to supply the soldiers, and urban residents with little or no production
of their own crops depended on large amounts of imported food supplies. This situation
meant not only more intensive agriculture in rural areas, but also an increasing importance
of markets and large-scale trade throughout the Mediterranean, with the frequent arrival
at Roman ports of cargoes from the islands of Sicily and Sardinia, North Africa and
Egypt [161,187,188].
To have cereals and legumes available throughout the year, it was necessary to store
them properly in specially designed structures, known as horrea, with a view to their long-
term conservation [
189
]. Latin authors such as Varro (Res Rusticae, 1st century BCE) and
Pliny the Elder (Naturalis Historia, 1st century CE) described different storage practices,
stating that the storage of raw cereals and legumes in a sealed, cool and dry environ-
ment would prevent germination and deterioration, allowing long-term storage. Under
conditions of high humidity and temperature, contamination and deterioration of the
entire supply would be encouraged, as well as rodent and insect infestations and mould
growth [
190
–
192
]. Rodents such as the wood mouse (Apodemus sylvaticus) (see Section 4.3.4.
Wild animals) were agricultural pests that damaged grain and seed stores [
95
,
153
]. Insects,
such as worms, weevils and beetles, could lay their eggs before cereals and legumes were
harvested, so larvae could emerge, feed on stored supplies, and contaminate them during
storage [
193
]. Mould could develop under ideal humidity and temperature conditions,
contaminating supplies by itself and attracting other insects that feed on the mould itself.
The horrea were not the only structures in Roman cities that stored, protected and
distributed cereals, legumes and other foods. Small outlets, such as bars, shops and
bakeries, represented important urban establishments where residents could buy these

Animals 2022,12, 213 21 of 37
foods or products made from them on a regular basis [
194
–
196
]. In cities such as Pompeii and
Herculaneum, various shops sold cereals, legumes, nuts and other foods [197]. To preserve
these food products, Roman shops and bars featured distinctive architectural elements,
such as large earthenware jars (dolia) embedded in a masonry service counter [
194
–
196
],
which provided them with great advantages. The ceramic material in these jars was an
effective thermal insulator capable of providing stable levels of temperature and humidity
for the contents. In addition, being enclosed in concrete counters, additional insulation was
provided to these containers. Thus, properly sealed, these jars protected their contents from
rodents, insects and other pests that could otherwise infiltrate. Therefore, this masonry retail
service counter worked in stores as a refrigerator for the preservation of food, especially
cereals and legumes, in stable, cool and dry environments [190].
Other structures of considerable interest are those reserved for the elaboration of
certain foods. The kitchens of most Roman houses did not have sufficient facilities for
making bread, so it was bought in the pistrinum (bakery). The pistor (baker) and his
assistants were in charge of the entire production process in the same establishment, which
had various specific rooms for each task: (i) storage: textile sacks of cereals were stored
in the attic of the establishment; (ii) grinding: from the attic, the sacks were poured onto
the catillus, the upper concave and hour-glass stone of the mill that rotates over the meta,
the stationary lower cone-shaped stone, both made of strong volcanic rock. Both pieces
formed a grinding wheel that was turned by animals or by humans—usually slaves—from
which the flour was produced. The animal-driven rotary mill significantly increased the
flour output and quality as equids (i.e., horses, donkeys or mules) could drive the mill for
hours at a time, saving the humans from the drudgery of milling; (iii) kneading: in another
area of the establishment there were kneading benches; (iv) baking: once the bread was
shaped, it was passed to the praefurnium (oven); and (v) sale: the establishment faced the
street, where the baked bread was taken for sale [198,199] (Figure 6a).
Animals 2022, 12, x 22 of 38
Figure 6. (a) The remains of the Pistrinum of Popidius Priscus (Pompeii) is a fine example of a bakery
in which the whole cycle of breadmaking from milling to baking the bread was performed. The
pistrinum contains four large millstones made from porous lava, traces of a stable, four storage
rooms and a large oven, which was used for baking the bread [Author: Carole Raddato. Distributed
under a CC BY-SA 2.0 license. Available online: https://cutt.ly/XRCZVTW. Accessed: 11 August
2021. (b) Thermopolium in Herculaneum, a commercial establishment where it was possible to
purchase ready-to-eat food [Author: Carole Raddato. Distributed under a CC BY-SA 2.0 license.
Available online: https://cutt.ly/FRCZMgT. Accessed: 30 October 2021]. All images are of public
domain via Wikimedia Commons webpage.
The large presence of ovens and pastry shops for the production and sale of flour-
based products suggests that baking was one of the most flourishing activities in Roman
cities [199]. The larger establishments were almost always equipped with grinding
wheels, a stable (the animals worked both in the milling of the cereal and in the transport
of supplies and processed products when necessary) and in a residential area, but without
a direct sales shop. The smaller establishments were more specifically dedicated to the
sale of bread (pistrina, bakeries) and pastry (pistrina dulciaria, pastry-shops) [200].
Thirty-five pistrina have been identified in Pompeii (e.g., House of the Chaste Lovers,
Bakery of Popidius Priscus) and another two in Herculaneum (e.g., Bakery of Sextus Patulcius
Felix) [172,201], some of them equipped with stables where skeletal remains of equines
have been found. Interesting is the House of the Chaste Lovers in Pompeii, the property of a
rich baker named Caius Iulius Polybius. This house had an adjoining bakery with a
splendidly preserved oven, millstones and two stables with the skeletal remains of five
equids. Adjacent to one of the stables, a little flat with a closet was found, which was
probably inhabited by the groom. The stable boy was in charge of cleaning the stable, the
equids, giving them food, changing the bedding and all other activities connected with
the stable [124].
In this way, working and consuming contaminated and deteriorated legumes and
cereals entailed health problems for millers, bakers and consumers [193], as testified in
the ancient sources as well as in the clinical findings, diagnoses and cures of many of the
diseases described in medical treatises, such as Galen’s (De Differentiis Febrium, De
Alimentorum Facultatibus; 2nd century CE) [191]. Warehouse workers, millers, bakers and
others who handled grain were vulnerable to skin reactions, inhalant allergies and
pathogens. The use of low-quality grain was also problematic, not only because it was less
nutritious, but also because the consumption of bread made with spoiled grain could lead
to health problems since pathogens were easily transmitted when people ingested insects
or moulds that contaminated them [190,191]. In addition, the stable, mill and bakery were
usually connected because they were part of the same bread-making process, so the close
contact of humans with equids propitiated the ideal environment to acquire infections—
parasitic, bacterial, viral and fungal—that could be transmitted by direct contact with
Figure 6.
(
a
) The remains of the Pistrinum of Popidius Priscus (Pompeii) is a fine example of a bakery
in which the whole cycle of breadmaking from milling to baking the bread was performed. The
pistrinum contains four large millstones made from porous lava, traces of a stable, four storage rooms
and a large oven, which was used for baking the bread [Author: Carole Raddato. Distributed under
a CC BY-SA 2.0 license. Available online: https://cutt.ly/XRCZVTW. Accessed: 11 August 2021.
(
b
)Thermopolium in Herculaneum, a commercial establishment where it was possible to purchase
ready-to-eat food [Author: Carole Raddato. Distributed under a CC BY-SA 2.0 license. Available
online: https://cutt.ly/FRCZMgT. Accessed: 30 October 2021]. All images are of public domain via
Wikimedia Commons webpage.
The large presence of ovens and pastry shops for the production and sale of flour-
based products suggests that baking was one of the most flourishing activities in Roman

Animals 2022,12, 213 22 of 37
cities [
199
]. The larger establishments were almost always equipped with grinding wheels,
a stable (the animals worked both in the milling of the cereal and in the transport of supplies
and processed products when necessary) and in a residential area, but without a direct
sales shop. The smaller establishments were more specifically dedicated to the sale of bread
(pistrina, bakeries) and pastry (pistrina dulciaria, pastry-shops) [200].
Thirty-five pistrina have been identified in Pompeii (e.g., House of the Chaste Lovers,
Bakery of Popidius Priscus) and another two in Herculaneum (e.g., Bakery of Sextus Patulcius
Felix) [
172
,
201
], some of them equipped with stables where skeletal remains of equines
have been found. Interesting is the House of the Chaste Lovers in Pompeii, the property
of a rich baker named Caius Iulius Polybius. This house had an adjoining bakery with a
splendidly preserved oven, millstones and two stables with the skeletal remains of five
equids. Adjacent to one of the stables, a little flat with a closet was found, which was
probably inhabited by the groom. The stable boy was in charge of cleaning the stable, the
equids, giving them food, changing the bedding and all other activities connected with the
stable [124].
In this way, working and consuming contaminated and deteriorated legumes and
cereals entailed health problems for millers, bakers and consumers [
193
], as testified in
the ancient sources as well as in the clinical findings, diagnoses and cures of many of
the diseases described in medical treatises, such as Galen’s (De Differentiis Febrium,De
Alimentorum Facultatibus; 2nd century CE) [
191
]. Warehouse workers, millers, bakers
and others who handled grain were vulnerable to skin reactions, inhalant allergies and
pathogens. The use of low-quality grain was also problematic, not only because it was less
nutritious, but also because the consumption of bread made with spoiled grain could lead
to health problems since pathogens were easily transmitted when people ingested insects
or moulds that contaminated them [
190
,
191
]. In addition, the stable, mill and bakery were
usually connected because they were part of the same bread-making process, so the close
contact of humans with equids propitiated the ideal environment to acquire infections—
parasitic, bacterial, viral and fungal—that could be transmitted by direct contact with these
animals, their urine or excrement or by the consumption of legumes, cereals and bread
that had been contaminated by coming into contact with the urine and excrement of these
animals during handling and processing.
4.4.5. Foodborne Zoonotic Diseases in Taverns (Thermopolia,Cauponae,Popinae) and Home
The complex theme of the diet in Roman times constituted and continues to present
a fertile area of research, also in relation to the state of health and disease of ancient
Roman populations. In fact, from the perspective of studies increasingly focused on a
multidisciplinary approach, and therefore on cooperation between different disciplines such
as archaeology, archaeozoology, archaeobotany, physical anthropology, classical literature
and the history of Roman art [
73
], an attempt is being made to outline an ever more
comprehensive picture both regarding the complex variety of food products available
on the market and consumed by at least part of the Roman population, as well as the
methods of preparation, handling and contamination of food. In particular, the issue of
food contamination by pathogens is of considerable interest as it allows us to advance
hypotheses about the state of health and disease of ancient populations in close relationship
with eating habits. After all, it is not surprising how, in ancient times, the populations
could suffer from different diseases associated with the consumption of contaminated food,
as it is known that some foods were not subjected to adequate processing before their
consumption and therefore could become possible vehicles for the transmission of zoonotic
diseases. The sources from which to outline the eating habits of the Vesuvian region are
numerous. Latin writers like Apicius (De Re Coquinaria, 1st century CE), Columella (De Re
Rustica, 1st century CE) and Pliny the Elder (Naturalis Historia, 1st century CE) talk about
Pompeian cooking. Their works offer a comprehensive idea of what should not be missing
in the kitchens and tables of the people of Pompeii. Other evidence of priceless historical
value comes from Herculaneum,Oplontis and Stabiae. These sites have yielded frescoes and

Animals 2022,12, 213 23 of 37
mosaics showing convivial scenes, remains and residues of crops and charred remains of
food in perfect condition of preservation.
No doubt, the culinary habits of the ancient Roman world deserve a more extensive
approach from the sanitary point of view, since animals destined for food are the main
reservoirs of many zoonotic foodborne pathogens, and food products of animal origin
are the main transmission vehicles [
12
]. Foodborne zoonotic diseases are caused by the
consumption of food contaminated by pathogenic microorganisms such as bacteria, viruses
and parasites. The risks of contamination can occur at any point along the food chain: at
the farm, during the slaughter and during processing or preparation and distribution [
12
].
It can also occur in taverns (e.g., thermopolia,cauponae,popinae) or in the home, if food is
incorrectly handled or cooked (Figure 6b).
From Pompeii and Herculaneum comes a large quantity of zooarchaeological material
associated with slaughtered bones and/or with burn marks that constitute tangible proof
that meat was most frequently consumed on Roman tables, as well as providing informa-
tion regarding the methods of handling (e.g., slaughter) and preparation (e.g., cooking).
Although later than the time of the eruption, of notable interest is the Edictum de Pretiis
Rerum Venalium (Edict Concerning the Sale Price of Goods), also known as Diocletian’s Edict
on Maximum Prices, which was an edict promulgated by the Roman Emperor Diocletian
that set maximum prices for more than 1300 products, as well as to establish labour costs
to produce them [
202
]. According to the Diocletian’s Edict, the most affordable protein
included sea fish, river fish, eggs, sheep’s milk and fresh cheese [202,203].
Meat sources included sheep/goat, beef, pork and poultry. In addition to these farm
animals, the Romans also ate wild animals, such as wild boar, deer, wild donkeys, chamois,
hare and birds, and other animals considered exotic such as flamingos and dormice [
204
]
(see Section 4.3.2. Livestock for more details on the dormouse). However, the popular
farcimina (spelt and meat sausages) were the cheapest meat and probably the unhealthiest
since they combined low-quality ingredients with intense handling [81].
The excavation of an ancient sewer in the city of Herculaneum provided an opportunity
to study the Roman diet in the Gulf of Naples, including marine resources used as food.
The remains of human and kitchen waste that accumulated in the sewer—representing
the waste of middle- and lower-class Romans that lived in an urban context—showed a
high degree of dietary diversity, with around 70 fish, 48 marine molluscs and 3 identified
marine arthropod taxa [
64
]. With regard to fish, the highest quantities found correspond to
black seabream (Spondyliosoma cantharus), damselfish (Chromis chromis), followed by seven
identified Sparidae species. Of the edible shellfish, the only species recovered in extremely
large quantities were the corneous wedge clam (Donacilla cornea), limpet (Patella spp.) and
purple sea urchin (Paracentrotus lividus) [
205
]. Both fish and shellfish would have been
available locally in large quantities and probably represented cheap and nutritious dietary
fare. Some of the fish consumed was likely salted (salsamenta) or fermented (such as fish
sauces such as garum), but it is believed that most of the fish and shellfish originated in the
Gulf of Naples and would have been purchased fresh [64].
Among all the foods that the Romans consumed and appreciated in a particular way,
and which may have represented a food vehicle for the infection and transmission of
intestinal parasites, there is certainly the garum [
206
]. The consumption and production of
garum, a sauce obtained from the fermentation in brine of whole fish products or parts of
them, including entrails and blood mixed with different spices, wine and vinegar, was also
very frequent in Herculaneum and Pompeii [
207
]. Much of the evidence about this ancient fish
sauce comes from classical literary sources (e.g., Pliny the Elder Naturalis Historia XXI.93ff,
1st century CE; Apicius De Re Coquinaria, 1st century CE). Furthermore, Pliny the Elder
(Naturalis Historia XXI.94; 1st century CE) cites Pompeii, together with Betica,Mauretania
Tingitana and Leptis Magna, as one of the most famous places of production and export of
garum in the Mediterranean. Likely, Pompeii had great fortune in the garum production
industry thanks to its privileged position in the Gulf of Naples and near the mouth of the
Sarno river, where the supply of salt together with a wide range of fish resources must

Animals 2022,12, 213 24 of 37
certainly have been abundant [
207
]. Generally, the processing of the fish used to obtain
garum by salting took place in special production areas located in suburban areas, within
which innumerable tubs or containers were used for maceration, as evidenced by the
discovery of an industrial area linked to the processing of fish products near Porta Stabia
at Pompeii [
208
]. The procedure for producing this famous sauce involved sun-drying fish
between 2 and 3 months; however, there were cases in which maceration was accelerated
by exposure to other heat sources such as ovens [
209
]. In Pompeii, the recent discovery of a
‘garum shop’, a building interpreted as a place of production of this sauce, not only allowed
more in-depth zooarchaeological analyses to be carried out on the remains of the fish found
still in a state of maceration in six perfectly preserved dolia at the time of the 79 CE eruption,
but also the obtention of more information about the production techniques [210–212].
Both meat and fish infected or contaminated during their handling and preparation
for consumption could transmit zoonotic diseases like those we see today if they were
consumed undercooked, subjected to mild brines, or lightly smoked. Thus, diseases caused
by Mycobacterium spp., Campylobacter spp., Salmonella spp., Streptococcus iniae,Clostridium
botulinum,Listeria monocytogenes or Vibrio vulnificus, and some parasites, such as Cryp-
tosporidium spp., Balantidium spp., Taenia saginata,Giardia lamblia or Toxoplasma gondii,
would not have been rare [
2
,
12
,
213
–
215
]. Hippocrates (Corpus Hippocraticum, Book of Epi-
demics; 4th century BCE) even recommended cooking meat and seafood well to avoid
diarrhoea [216].
Foodborne zoonoses acquired from meat are too many to mention, but perhaps bovine
tuberculosis could be considered a probable zoonotic disease in Roman times. In fact,
pathological skeletal changes to the ribs and vertebrae indicative of tuberculosis have been
documented in two adults in the Herculaneum osteological sample [
217
]. Tuberculosis
is caused by a bacterium that belongs to the genus Mycobacterium, and Capasso and Di
Tota [
217
] suggested that it was possible that the disease may have been contracted by the
consumption of undercooked infected oxen viscera after ritual sacrifice, though there were
probably other sources of infection.
There are several literary sources that testify that in Roman times sheep farming was
frequently cited not only in relation to the consumption of meat, but also to the production
of wool, milk and its derivates (Varro Res Rusticae II.8, II.11.1, 1st century BCE; Columella
De Re Rustica VII.2, 1st century CE; Pliny the Elder Naturalis Historia VIII.123–124, 180, 1st
century CE). With regard to sheep’s milk, the Romans considered it an essential food in
their daily diet (Varro Res Rusticae II.2.2, 1st century BCE; Pliny the Elder Naturalis Historia
XXVIII, XXXIII, CXXIV, 1st century CE), so much so that even Celsus in his medical treatise
recommended its intake, underlining its health benefits (Celsus De Medicina, 1st century
CE). Particularly appreciated, as well as being considered a particularly delicious food,
was the colostrum (the milk of a sheep that had just given birth). Further information
and confirmation about the use of this food can be found in the work of Apicius (De Re
Coquinaria VII, XI, XIII, 1st century CE). In his corpus of recipes, the author, among the
various products handed down in the culinary field, mentions milk as a fundamental basis
for the preparation of desserts, creams, cheeses and sauces. Speaking of sauces, the Romans
used to consume melca, a kind of yoghurt or sour cream obtained by using sour milk mixed
with different spices, such as pepper, salt, oil, coriander, old-fashioned melca, herbs and
onions [
218
,
219
]. More generally, fresh and unpasteurized raw milk, mixed with fig juice
regurgitated by young lambs not yet weaned, was used to produce fresh cheeses (Pliny
the Elder Naturalis Historia XI.239, 1st century CE). Fresh cheeses were usually intended
mostly for quick consumption, unlike those that were seasoned with the aid of salt. In this
case, according to what is always learned from literary sources (Varro Res Rusticae II.2.3, 1st
century BCE; Columella De Re Rustica VII.2.1, VII.8.1, 1st century CE), which accurately
describe the production procedures of fresh and aged cheeses, there were cases in which
dairy products were preserved by previous drying in the sun and subsequent pressing
and immersion in brine [
207
,
220
]. In central and southern Italy, the milk, if not used for

Animals 2022,12, 213 25 of 37
the preparation of cheeses, was eaten fresh during the day or curdled with the addition of
aromatic herbs given the difficulty of preservation.
Renowned in coeval sources for the production and consumption of sheep and goat
milk, the ancient Vesuvian city of Herculaneum has given us back several portions of charred
cheese, whose microbiological examination has allowed us to infer information about
how these foods were consumed and used in Roman gastronomy, as well as in relation
to their processing methods. In fact, inside one of these charred cheeses was found the
presence of spherical-shaped bacteria that, both in terms of morphology and dimensions,
seemed to be consistent with the genus Brucella. The heat of the eruption, coupled with
the subsequent burial environment, made it impossible to use molecular techniques to
confirm identification [
61
]. Moreover, Capasso [
221
] described vertebral lesions associated
with rib alterations in 16 adult individuals of the Herculaneum skeletal sample, which
suggested brucellosis. The high frequency of the disease (about 17.4% of the adults) is also
supported by historic evidence (Varro Res Rusticae; II.5.4, 1st century BCE; Cicero De Natura
Deorum II.159, 1st century BCE; Pliny the Elder Naturalis Historia VIII.180, 1st century CE)
and the carbonized remains of cheese. Thus, the historical and biological data from the
Herculaneum population provides the first evidence of a relationship between milk-related
sources and brucellosis.
In Roman cuisine, it was also common to use and consume other foods of animal
origin through which it was possible to contract some zoonotic diseases. Among these
were, for example, the eggs, not only of hens but also of goose, ducks, peacocks and wild
birds. Eggs were a widely frequent food in Roman gastronomy. Apicius (De Re Coquinaria,
1st century CE) reports their use in various recipes as a basis for the preparation of sauces,
binders and desserts. Among the countless organic remains found in Herculaneum, eggs
were also among the foods contaminated by pathogens that may have negatively affected
the health of the population. As in the case of cheese, the microbiological examination
performed on the interior surface of the preserved charred eggshell of hard-boiled eggs
from Herculaneum also showed the presence of a cocciform bacterium morphologically
and dimensionally consistent with the genus Salmonella [
62
]. In the past, as in modern
times, salmonellosis could be transmitted to humans via the faecal—oral route, that is,
either through the ingestion of contaminated foods, such as in the case of eggs, or through
the contact and/or handling of objects soiled by the faeces of small domestic animals
affected by Salmonella [
78
,
222
]. Therefore, infections in ancient Roman populations could be
contracted both during the collection of the eggs, such as through the use and manipulation
of these during the preparation of some dishes that required their use [
222
]. Albeit in a
risky way, even in the case of eggs, it can be assumed that these could have constituted a
food that could have acted as a vehicle for the diffusion of zoonosis in humans [78].
4.5. Latrines, Sewers and Baths (Thermae)
In some houses or in certain corners of the insulae could be found sterquilinia (private
latrines), located next to the kitchen or even inside it. This disposition was understood
to easily get rid of organic waste and excess liquids since the number of pipes in the
houses was reduced, thus simplifying and making their construction cheaper; however,
the risk of food contamination in such combined kitchen/latrine areas must have been
very high [
85
]. In addition, these latrines were not usually cleaned with water, nor were
they connected to the municipal sewer system, but instead led directly to cesspits built
with porous stone and without mortar, which allowed the drainage of liquids, leaving
the solid residue to be cleaned periodically [
85
,
200
]. It is believed that this disconnection
of the latrines from the sewer system could be due to avoiding possible backflows and
accumulations of gases, evading the mandatory payment of the right to use the sewage
system or preventing the entry of animals and insects from the sewers. However, it seems
clear that the most likely cause was the value that faeces had as manure from both animal
and human waste [
167
]. According to the written sources (Martial Epigrammata I.37.1–2,
1st century CE; Petronius Satyricon XXVII.3, XLI.9, XLVII.5, 1st century CE), urinals of

Animals 2022,12, 213 26 of 37
various types also abounded, such as the matellae and the lasana—vessels for the reception
of human urine and excrements, which the slaves (known as lasanophoroi) emptied into the
latrines [85,223].
On the other hand, the foricae (communal latrines) seemed to be a flourishing business,
which had jugs located in the streets where, for a modest tax, it was possible to urinate [
85
].
Lucretius (De Rerum Natura 4.1026–1029, 1st century BCE) refers to the use of vessels named
dolia curta as a urinal/urine container—these containers were terracotta amphorae with their
upper portion removed [
179
,
224
]. The foricarii and stercorarii were in charge of keeping the
latrines, sewers and urinals clean, as well as transporting in a plaustrum—an open wagon
or cart for hauling loads—the waste from the city to rural districts, where they were sold to
farmers as manure [
162
]. Columella (De Re Rustica I.6.24, 1st century CE) already mentions
that human faeces were recycled into fertilizer for crops (see Section 4.4.1. Agriculture), so
that an entire infrastructure had been developed for its use. In addition, urine was used as
a detergent in the fullonicae, establishments dedicated to the cleaning of fabrics and wool;
see Section 4.4.3. Textile industry: Laundries (fullonicae): as a mordant for some dyes in
the officinae infectoriae, workshops specializing in the dyeing of new textile products; in the
officinae offectoriae, workshops for the dyeing of used textile products; and as a bleach in
officinae tinctoriae [
173
]. Hence, the installation of dolia curta on public roads to be able to
collect it [224].
Cesspits retained their popularity into the 2nd century CE, well after public toilets,
flushed by sewer drains, were firmly established in the urban fabric. The persistence of
cesspits probably had as much to do with simple economics than with resistance to change.
A city notice found at Herculaneum (exemta ste(r)cora a(ssibus) XI) illustrates that 11 donkeys
might have been paid for the removal of manure [
74
] (p. 101). Foricarii and stercorarii
cleared it from the streets but also emptied cesspits and sold the contents to farmers on the
outskirts of the city. The 11 donkeys mentioned in the Herculaneum inscription was not a
very substantial sum, but when that sum was added to a farmer’s payment for the manure,
it can be seen as being of modest economic value for the poorer urban dweller who was
willing to take on this odious task [85].
The problems associated with the disposal of human waste could be of a different order
or nature, starting from the emanation of bad smells that acted as an attractive element for
some animals, which, coming into contact with infected material, became, in turn, a vehicle
for the transmission of pathogens moving freely around the city [
85
,
167
]. Generally, in
Roman cities, the disposal of excrement and waste was closely linked to a complex system
of public sewers, both public (foricae) and private (sterquilinia) latrines or simple cesspits
located near the house that facilitated the unloading and drainage of wastewater. Despite
the rigorous norms of Roman sanitary hygiene legislation, it was not mandatory to be
connected to a public sewage system since this was a paid service [
85
]. For the houses
or shops that were not connected, there was a private service in which the foricarii and
stercorarii—generally people of lower social status, mostly prisoners and slaves—came to
the building every day to collect waste from clay containers, the content of which was sold
to farmers outside the city [
85
,
162
,
163
]. However, the conditions varied throughout the
Empire. For example, in Pompeii, buckets containing waste and faecal matter were thrown
out of the windows directly onto the street below, where they accumulated and dispersed,
favouring the spread of diseases. The installation of stepping-stones documents this fact so
that people could cross the street without getting their feet wet [
225
] (Figure 7a). To keep
the streets clean, the aediles—the town authorities—of Herculaneum established a series of
prohibitions and orders, such as a ban on throwing garbage on the ground near public
drinking fountains. In an information panel located directly near the fountain, the aediles
established sanctions for this violation: citizens had to pay a fine, and slaves had to be
flogged [
225
,
226
]. However, even the enviable sewers did not guarantee complete health
since their maintenance could be insufficient. For example, Caligula punished Vespasian—
still a mere mayor—for not keeping the streets and sewers clean, and Julius Caesar forced
citizens to take care of cleaning their streets [225]. In addition, foricarii and stercorarii were

Animals 2022,12, 213 27 of 37
exposed to rats, insects and wastewaters, which frequently stagnated, and at times the
same rainwater filled the sewers and cloacae and carried excrement and debris to the
surface [
85
]. It is thus clear that there were numerous health implications as a consequence
of the reuse of human faeces and urine, such as (i) plants fertilized with human—and
animal—manure and converted into a probable source of pathogens; (ii) losses during
transport in porous containers—such as the dolia curta, which were made of terracotta—and
in the plaustra—obviously these carts were not watertight; (iii) handling of urine and faeces
by foricarii and stercorarii; (iv) dunghills and the accumulation of faeces and urine prior to
transportation; (v) contamination of waterpipes; or (vi) presence of puddles in the absence
of adequate drainage systems [85].
Animals 2022, 12, x 28 of 38
Figure 7. (a) Stepping-stones in Via Stabiana (Pompeii), whose function was to keep Romans’ feet dry
and out of the rainwater, slop and animal waste that would have filled the streets [Author: Jensens.
Distributed under a CC0 license. Available online: https://cutt.ly/aRCZ8V8. Accessed: 30 October
2021]. (b) Reconstruction drawing showing the communal latrines in use [Author: Carole Raddato.
Distributed under a CC BY-SA 2.0 license. Available online: https://cutt.ly/dRCZ471. Accessed: 30
October 2021]. (c) Replica of xylospongium (sponge on a stick). It was a hygienic tool used and shared
in communal latrines by ancient Romans to wipe their buttocks after defecating [Author: D.
Herdemerten. Distributed under a CC BY 3.0 license. Available online: https://cutt.ly/iRCZ7GO.
Accessed: 30 October 2021]. All images are of public domain via Wikimedia Commons webpage.
More than 88 latrines have been discovered in the portion of Herculaneum that has
been excavated. Out of 41 houses, 28 of them had one or more latrines, producing a total
of 39. Of these, 14 were located next to a kitchen, while the rest were set in the servants’
quarters or service rooms. Latrines have also been found in many shops (tabernae) and
workshops. Of the 68 commercial premises at Herculaneum, 27 had a toilet near the
entrance, which might have benefitted not just the workers but also their clients [227]. In
the case of latrines inside the kitchens or tabernae (shops selling foodstuffs), poor
management of human waste disposal could pose a significant risk of accidental
contamination of food with pathogens [75,77,170]. Finally, although the forum of
Herculaneum has never been excavated and its various public buildings have only been
partially uncovered, 11 public buildings were discovered, in which 9 latrines have been
found [227]. It is interesting to note that the foricae (public latrines) were sometimes
attached to thermae (baths) [162]. In the case of Herculaneum, four of the public latrines
were in the Central Baths and one in the Suburban Baths [227].
The use of thermae and public latrines was a very widespread and common practice
in the Roman custom that the entire population enjoyed daily. In particular, the thermae
did not only serve as sanitation facilities but were also a place for social gathering and
conversation [167] (Figure 7b). In the case of public latrines, these represented an
environment in which it was possible to come into contact with parasites that could have
negative health consequences on people’s health. The xylospongium or tersorium—also
known by the sources as a hygienic tool used and shared in public latrines by the
Romans—was a sponge fixed to a stick used to clean the buttocks after defecation and
may well have been one of the vehicles responsible for the spread of parasites and
intestinal tract bacteria in humans [167,228–230] (Figure 7c). Both Seneca (Epistulae morales
ad Lucilium VIII.70.20, 1st century CE) and Martial (Epigrammata XII.48.7, 1st century CE),
even if they do not describe the method of use, imply the context of use, which denotes
the use of xylospongium in ancient latrines [85]. Moreover, widespread in the Roman world
could have been the custom of not following strict hygiene rules regarding hand washing
after using the latrines and after preparing food; in this case, this practice would suggest
Figure 7.
(
a
) Stepping-stones in Via Stabiana (Pompeii), whose function was to keep Romans’ feet dry
and out of the rainwater, slop and animal waste that would have filled the streets [Author: Jensens.
Distributed under a CC0 license. Available online: https://cutt.ly/aRCZ8V8. Accessed: 30 October
2021]. (
b
) Reconstruction drawing showing the communal latrines in use [Author: Carole Raddato.
Distributed under a CC BY-SA 2.0 license. Available online: https://cutt.ly/dRCZ471. Accessed:
30 October 2021]. (
c
) Replica of xylospongium (sponge on a stick). It was a hygienic tool used and
shared in communal latrines by ancient Romans to wipe their buttocks after defecating [Author: D.
Herdemerten. Distributed under a CC BY 3.0 license. Available online: https://cutt.ly/iRCZ7GO.
Accessed: 30 October 2021]. All images are of public domain via Wikimedia Commons webpage.
More than 88 latrines have been discovered in the portion of Herculaneum that has
been excavated. Out of 41 houses, 28 of them had one or more latrines, producing a total
of 39. Of these, 14 were located next to a kitchen, while the rest were set in the servants’
quarters or service rooms. Latrines have also been found in many shops (tabernae) and
workshops. Of the 68 commercial premises at Herculaneum, 27 had a toilet near the entrance,
which might have benefitted not just the workers but also their clients [
227
]. In the case
of latrines inside the kitchens or tabernae (shops selling foodstuffs), poor management of
human waste disposal could pose a significant risk of accidental contamination of food
with pathogens [
75
,
77
,
170
]. Finally, although the forum of Herculaneum has never been
excavated and its various public buildings have only been partially uncovered, 11 public
buildings were discovered, in which 9 latrines have been found [
227
]. It is interesting to
note that the foricae (public latrines) were sometimes attached to thermae (baths) [
162
]. In
the case of Herculaneum, four of the public latrines were in the Central Baths and one in the
Suburban Baths [227].
The use of thermae and public latrines was a very widespread and common practice
in the Roman custom that the entire population enjoyed daily. In particular, the thermae
did not only serve as sanitation facilities but were also a place for social gathering and
conversation [
167
] (Figure 7b). In the case of public latrines, these represented an environ-

Animals 2022,12, 213 28 of 37
ment in which it was possible to come into contact with parasites that could have negative
health consequences on people’s health. The xylospongium or tersorium—also known by
the sources as a hygienic tool used and shared in public latrines by the Romans—was a
sponge fixed to a stick used to clean the buttocks after defecation and may well have been
one of the vehicles responsible for the spread of parasites and intestinal tract bacteria in
humans [
167
,
228
–
230
] (Figure 7c). Both Seneca (Epistulae morales ad Lucilium VIII.70.20,
1st century CE) and Martial (Epigrammata XII.48.7, 1st century CE), even if they do not
describe the method of use, imply the context of use, which denotes the use of xylospongium
in ancient latrines [
85
]. Moreover, widespread in the Roman world could have been the
custom of not following strict hygiene rules regarding hand washing after using the latrines
and after preparing food; in this case, this practice would suggest a further form of contam-
ination of the food that was then ingested. This is especially the case in the consumption
of fruit and vegetables, which did not always require a specific cooking method useful
for killing any pathogens [
167
]. Like latrines, the thermae (baths) would have represented
another environment potentially suitable for the proliferation and spread of zoonotic dis-
eases because some zoonoses are also waterborne [
231
]. In the case of the thermae, it is
possible that the water was not changed regularly and that consequently, this aspect led
to an accumulation of waste and at the same time created, through the stagnation of dirty
water and heat, an environment particularly suitable for the proliferation of parasites, such
as roundworms and protozoa, which in turn infected humans causing gastrointestinal
disorders, but some other vectorial diseases (viral and bacterial), including zoonoses like
leishmaniosis could have been present.
In order to better understand the close relationship between dietary diversity, strategies
for waste disposal and problems related to the public health of the population, once again
is interesting the analysis of biological material from the Cardo V sewer at Herculaneum.
This biological material includes human excrement and the remains of edible foodstuffs,
including eggshells, seashells, fish bones, otoliths and botanical material. In general, this
sewer lacked an outflow point and instead functioned as a cesspit to collect the human and
kitchen waste generated by those that lived in a mixed commercial and residential structure
whose shops and apartments were situated above (in the Insula Orientalis II) [
64
,
65
,
205
].
Several studies have been carried out on the faecal material coming from the latrines of
Herculaneum, which confirm the presence and diffusion of both various intestinal parasites
and of their eggs due to the contamination of food and water by human faeces; among
the most common predominate the whipworm (Trichuris trichiura),roundworm (Ascaris
lumbricoides) and Entamoeba histolytica, which causes dysentery [
75
,
206
,
232
]. In a recent
study, Ledger et al. [
233
] also observed helminth eggs of Trichuris trichiura and Ascaris
lumbricoides when analysing pelvic sediment samples from skeletal remains from Oplontis.
All these aspects would suggest how, despite the presence and use of public latrines being
quite widespread in the Roman Empire, together with a complex system of washing plants,
sewage networks and drinking water conveyed by aqueducts within the city, and even
though hygiene rules and the disposal of human waste were not neglected, still these
measures were insufficient to protect the population from the possibility of contracting
parasites spread through faecal contamination.
4.6. Rituals
For the ancient Romans, animal sacrifice was performed as a ritual to communicate
with gods, heroes and other divine beings. Such rituals were intended to request favours,
protection and help from divine recipients or to appease them through prayer. The sacrifice
of animals was also a way for human worshippers to know the will of the gods, and the
ritual often concluded with the distribution and consumption of the meat, which served to
strengthen and define the social fabric by marking who belonged to a particular group and
who was an outsider, expressed largely by the degree of access to meat [
234
]. The main
evidence available for the study of ancient animal sacrifice is provided by literary texts,
inscriptions, frescoes and archaeological remains in the form of altars and other sacrificial

Animals 2022,12, 213 29 of 37
facilities, as well as animal skeletal remains [
235
–
238
]. These lines of evidence show that
the animals chosen for sacrifice were generally of domestic species, such as cattle, sheep,
goats or pigs.
In consideration of the use of livestock animals for ritual purposes, it is possible to
hypothesize how this practice could have represented a further channel of transmission
of zoonotic diseases. In fact, following the sacrifices, the animals were eviscerated pri-
marily to make the offering to the gods (e.g., liver, heart, lungs, spleen, intestines) and
subsequently to examine the contents of the internal organs to interpret the liking of the
gods themselves [
234
]. Following the public sacrifice, the meat was distributed and served
among those who had taken part in the sacrifice itself, and in some cases, to the population.
In order for the worshippers to be able to consume the meat, the viscera and other remains
of the animal, they first had to be returned to the profane sphere—in order to deprive
them of a certain sacredness—which was done by boiling the meat in milk or simply in
water, or the sacrificer placing his hand on the carcass, a gesture that transformed the meat
into something that men could eat. From this moment, the meat could be divided and
distributed [
234
]. The meat was often eaten in the sanctuary where the sacrifice had been,
but it could also be carried in small sportulae—food baskets distributed as gifts by wealthy
and influential people to clients and protégés [
87
] (p. 209)—to consume at home or sell in
the macella—public markets dedicated to the sale of meat and fish [239].
It is evident that sacrificial meat was an important source of protein in the diet in
ancient times, and meat consumed at public banquets appears to be derived from ritual
sacrifices [
240
]. In the Roman macella, wild birds (e.g., pheasants and pigeons) and fish were
sold, in addition to the most common meat of beef, lamb and pork [
241
,
242
]. Recent studies
of the macella have shown that these facilities offered meat from both shrine sacrifices and
from animals sacrificed in the market building, which was equipped with altars and statues
of deities [
239
]. This situation is very evident in Pompeii, where the macellum is in the centre
of the forum, which allows easy access to this open space, and in the vicinity of which there
were a series of tabernae (shops selling foodstuffs) and more than ten temples. The animals
sacrificed in these sanctuaries could have been sacrificed and sold in the macellum, as seems
to be evidenced by the discovery of a small facility containing the bones of animals that
died during the eruption of Vesuvius [234,243] (p. 467).
5. Concluding Remarks
There is no doubt that the cultural and urban environments contributed to the animal–
human interaction in the daily life of the ancient Roman cities of Pompeii and Herculaneum.
As testified in ancient medical (e.g., Celsus’ De Medicina; Galen’s volumes) and veterinary
(e.g., Vegetius’ Digesta Artis Mulomedicinalis) treatises, numerous diagnoses and cures in re-
lation to zoonotic diseases were outlined. Summarising, the main drivers and mechanisms
for the distribution and transmission of zoonotic diseases could have been: (i) the large
number of and the role played by different animal species in the ancient Roman world
(e.g., livestock, wild, exotic and household animals); (ii) the environmental conditions for
the survival of parasites, pathogens and vectors (e.g., population density, substandard
housing, different available materials for the construction of the houses, use of animal and
human wastes as fertilizer, lack of cleaning and hygiene); (iii) the great variety and intensity
of commercial activities and occupations that represented certain risks of infections due
to the continuous contact with animals and their products (e.g., agriculture and farming,
textile industry, laundries, bakeries); (iv) the absence of adequate safety controls during the
processing, distribution and preservation of foodstuffs in unsuitable environments, and
some culinary habits (e.g., incorrect handling and cooking); (v) the inadequate mechanisms
of the disposal of human waste and the biotic contamination of watercourses and reservoirs
(e.g., combined kitchen/latrine areas, shared use of baths and public latrines); and (vi) the
use of animals related to common cultural and religious practices (e.g., participation in
public entertainment games, sacrifice as offerings to the gods).

Animals 2022,12, 213 30 of 37
Author Contributions:
Conceptualization, J.V.; investigation, C.T., M.R. and J.V.; writing—original
draft preparation, C.T., M.R. and J.V.; writing—review and editing, C.T., M.R. and J.V.; supervision,
J.V. All authors have read and agreed to the published version of the manuscript.
Funding: This research received no external funding.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Not applicable.
Conflicts of Interest: The authors declare no conflict of interest.
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