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medicine and biomedical sciences in modern history
ANIMALS AND THE SHAPING
OF MODERN MEDICINE
ONE HEALTH AND ITS HISTORIES
ABIGAIL WOODS, MICHAEL BRESALIER,

ANGELA CASSIDY, RACHEL MASON DENTINGER
Medicine and Biomedical Sciences
in Modern History
Series Editors
Carsten Timmermann
University of Manchester
Manchester
United Kingdom
Michael Worboys
University of Manchester
Manchester
United Kingdom
The aim of this series is to illuminate the development and impact of
medicine and the biomedical sciences in the modern era. The series was
founded by the late Professor John Pickstone, and its ambitions reflect
his commitment to the integrated study of medicine, science and tech-
nology in their contexts. He repeatedly commented that it was a pity
that the foundation discipline of the field, for which he popularized the
acronym ‘HSTM’ (History of Science, Technology and Medicine) had
been the history of science rather than the history of medicine. His point
was that historians of science had too often focused just on scientific
ideas and institutions, while historians of medicine always had to con-
sider the understanding, management and meanings of diseases in their
socio-economic, cultural, technological and political contexts. In the
event, most of the books in the series dealt with medicine and the bio-
medical sciences, and the changed series title reflects this. However, as
the new editors we share Professor Pickstone’s enthusiasm for the inte-
grated study of medicine, science and technology, encouraging studies
on biomedical science, translational medicine, clinical practice, disease
histories, medical technologies, medical specialisms and health policies.
The books in this series will present medicine and biomedical science
as crucial features of modern culture, analysing their economic, social
and political aspects, while not neglecting their expert content and con-
text. Our authors investigate the uses and consequences of technical
knowledge, and how it shaped, and was shaped by, particular economic,
social and political structures. In re-launching the Series, we hope to
build on its strengths but extend its geographical range beyond Western
Europe and North America.
Medicine and Biomedical Sciences in Modern History is intended to
supply analysis and stimulate debate. All books are based on searching
historical study of topics which are important, not least because they cut
across conventional academic boundaries. They should appeal not just to
historians, nor just to medical practitioners, scientists and engineers, but
to all who are interested in the place of medicine and biomedical sciences
in modern history.
More information about this series at

http://www.springer.com/series/15183
Abigail Woods · Michael Bresalier
Angela Cassidy · Rachel Mason Dentinger
Animals and the

Shaping of Modern
Medicine
One Health and its Histories
Abigail Woods Angela Cassidy
Department of History Department of Politics
Kings College London University of Exeter
London, UK Exeter, UK
Michael Bresalier Rachel Mason Dentinger

Department of History University of Utah
Swansea University Salt Lake City, UT, USA
Swansea, UK
Medicine and Biomedical Sciences in Modern History

ISBN 978-3-319-64336-6 ISBN 978-3-319-64337-3 (eBook)
https://doi.org/10.1007/978-3-319-64337-3
Library of Congress Control Number: 2017948270
© The Editor(s) (if applicable) and The Author(s) 2018. This book is an open access publication.
Open Access This book is licensed under the terms of the Creative Commons Attribution
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use, sharing, adaptation, distribution and reproduction in any medium or format, as long as
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Neither the publisher nor the authors or the editors give a warranty, express or implied,
with respect to the material contained herein or for any errors or omissions that may have
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maps and institutional affiliations.
Cover credit: [Plate eight - Man, cow and sheep and explanatory text], pp. [unnumbered]-22,
in Hawkins, Benjamin Waterhouse / A comparative view of the human and animal frame
(1860), Image courtesy of The University of Wisconsin Libraries
Printed on acid-free paper
This Palgrave Macmillan imprint is published by Springer Nature

The registered company is Springer International Publishing AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
To the animals (human and non-human) who brought us together.
Preface
It is not long since the question ‘Where are the animals in medical
history?’ prompted yawning and shuffling of feet among scholars of that
discipline. While in the wider world the health agenda known as ‘One
Medicine’ or ‘One Health’ was gathering momentum by highlighting
the deeply interconnected nature of human and animal health and the
need for integrated approaches to it, with a few key exceptions, scholars
in medical history continued to believe that the only animals important
to medicine were human animals. Conference organizers asked if animals
belonged on medical history programmes; conference delegates voted
with their feet; and scholarly discussions proceeded largely in ignorance
of how animals and animal health had shaped—and been shaped by—the
history of human health, medicine and society.
This was the situation that inspired the programme of research on
which this volume is based. Generously sponsored by the Wellcome
Trust,1 it set out to explore the zoological foundations of human medi-
cine, to illuminate the history of animals in medicine, and to develop an
empirically grounded history of the recent movement for One Health.
Research began at Imperial College London in 2011 and terminated
at King’s College London in 2016. It was conducted by a team of four
scholars—Abigail Woods (principal investigator), Michael Bresalier,
1Programme Grant reference 092719/Z/10/A.
vii
viii Preface
Rachel Mason Dentinger and Angela Cassidy—who are the joint authors
of this volume.
Combining first degrees in veterinary medicine and the life sci-
ences, with scholarly careers that straddle the history and sociology of
veterinary medicine, human medicine and biology, we formed an inter-
disciplinary team well equipped to study the history of medicine as an
interdisciplinary, interspecies phenomenon. Each of us has worked on
discrete research projects that address a different aspect of this issue. This
volume presents findings from each project, in five sample chapters that
bear the authors’ names.
However, the work as a whole is a shared endeavour. It grew out of
our many meetings, in which we reviewed existing historical accounts
of animals and medicine, and worked together to develop a shared lan-
guage, conceptual apparatus and approach to studying their intercon-
nected histories. It aspires to greater cohesion and coherence than a
standard edited volume. It was also more difficult to write—more diffi-
cult, even, than a standard monograph in which only a single author has
to make decisions about arguments and narrative. We found few prec-
edents to guide us: team working is relatively new to the discipline of
history, and, judging by certain publishers’ responses to the notion of a
volume with four authors, it is equally unfamiliar to academic publishing.
Consequently, we have had to develop our working, writing and publish-
ing practices by trial and error. This has been a very time-consuming but
ultimately fulfilling experience. The mutual support and advice of col-
leagues has pushed our scholarship to a higher level, and enabled us to
work on a broader canvas than would have been possible otherwise.
We are very grateful to Palgrave for supporting our vision and helping
us to realize it. We hope that our readers—whether medical historians,
animal historians or participants in One Health today—will find this an
interesting and a thought-provoking volume. We also hope that it will
persuade our colleagues in medical history that without asking ‘Where
are the animals?’ and ‘What do they do?’, we cannot truly understand
what has constituted medicine in history or what it has become today.
Many people have contributed to the preparation of this volume.
Collectively, we wish to thank the Wellcome Trust for funding our
research, colleagues (especially Dr. Kathryn Schoefert) in the Centre
for the History of Science, Technology and Medicine at King’s College
London for their ongoing support, and the various audiences, review-
ers and expert advisors who have provided constructive feedback on our
Preface ix
findings in the course of the research programme. Abigail Woods would

also like to thank Department III at the Max Planck Institute for the
History of Science, Berlin, for hosting her during the spring term of
2017, which was a crucial writing-up phase. Angela Cassidy would like
to thank new colleagues at the University of Exeter for their support
during writing up, and the scientists and veterinarians she interviewed
as part of this research for their essential insights into One Health and
disciplinary politics in the twenty-first century. Rachel Mason Dentinger
would also like to thank new colleagues at the University of Utah for
providing support during the completion of this book. Michael Bresalier
would like to thank archivists at the Food and Agriculture Organization
(Fabio Ciccarello) and the World Health Organization (Reynald Erard)
for their support of the research for his chapter, his new colleagues at
Swansea University for embracing his work, and Abigail Woods for her
remarkable support in completing the job.
London, UK Abigail Woods

Swansea, UK Michael Bresalier
Exeter, UK Angela Cassidy
Salt Lake City, USA Rachel Mason Dentinger
Contents
1 Introduction: Centring Animals Within Medical History 1

1.1 Why Animals? 4
1.2 Writing Animal Histories 7
1.3 Animals in Medical History 11
1.4 One Health and its Histories 14
Bibliography 20
2 Doctors in the Zoo: Connecting Human and Animal

Health in British Zoological Gardens, c.1828–1890 27
Abigail Woods
2.1 Disease and Death in the Zoo 32
2.1.1 Public Health 35
2.1.2 Bedside Medicine 39
2.1.3 Hospital Medicine 43
2.2 Comparative Perspectives 46
2.2.1 The Pursuit of Comparative Pathology 49
2.2.2 Tuberculosis and Rickets 54
2.3 Conclusion 58
Bibliography 61
xi
xii Contents
3 From Coordinated Campaigns to Watertight

Compartments: Diseased Sheep and their Investigation
in Britain, c.1880–1920 71
Abigail Woods
3.1 Coordinated Campaigns 75
3.2 Research Reconfigurations 87
3.3 Watertight Compartments 97
3.4 Conclusion 103
Bibliography 107
4 From Healthy Cows to Healthy Humans: Integrated

Approaches to World Hunger, c.1930–1965 119
Michael Bresalier
4.1 Cows in Interwar Medicine and Agriculture 123
4.2 War and its Aftermath 130
4.3 Healthy Cows, Healthy Humans 136
4.4 Conclusion 147
Bibliography 150
5 The Parasitological Pursuit: Crossing Species and

Disciplinary Boundaries with Calvin W. Schwabe and
the Echinococcus Tapeworm, 1956–1975 161
Rachel Mason Dentinger
5.1 Pursuing Echinococcus in Beirut: The Parasitology
of Calvin W. Schwabe 165
5.2 Echinococcus Leaves the Laboratory: Schwabe’s
Parasitology at the Population Level 171
5.3 Following Echinococcus Across the Globe: Schwabe’s
Persistent Parasitology 176
5.4 Conclusion 184
Bibliography 187
6 Humans, Other Animals and ‘One Health’ in the Early

Twenty-First Century 193
Angela Cassidy
6.1 One Health or Many? 198
6.1.1 Calvin Schwabe and One Medicine 200
6.1.2 The Wildlife Conservation Society 205
Contents xiii
6.1.3 The Swiss Tropical and Public Health Institute 207

6.1.4 The One Health Initiative and Commission 209
6.2 The Animal Subjects of One Health 214
6.2.1 Animals in One Health Research Texts 215
6.2.2 Animals in One Health Imagery 219
6.3 Conclusion 225
Bibliography 227
7 Conclusion 237
Appendix: Annotated Bibliography of Animals

in the History of Medicine 247
Index 269
List of Figures
Fig. 6.1 The One Health umbrella 199

Fig. 6.2 Citations to Schwabe’s VMHH, 1964, 1969, 1984 201
Fig. 6.3 Growth of the One Health bandwagon 215
Fig. 6.4 Human–animal partnerships 219
Fig. 6.5 OH contextualised 221
xv
List of Tables
Table 3.1 British research into diseased sheep, 1880–1901 78

Table 3.2 British research into diseased sheep, 1902–1920 89
Table 6.1 One Health and animal categories 216
Table 6.2 One Health and animal species 217
xvii
CHAPTER 1
Introduction: Centring Animals

Within Medical History
In a recent handbook on the history of medicine, authors Robert Kirk

and Michael Worboys argued that ‘In no body of scholarship is it more
obvious, puzzling and true to say that “animals disappear.”’1 Literally, of
course, this is not the case, for as Etienne Benson points out,
to a limited but important extent, writing about human history is always—

already writing about animals … Humans are a kind of animal that (like all
kinds of animal) has been and continues to be profoundly reshaped by its
interactions with other kinds of animals … All history is animal history, in
a sense.
However, Benson acknowledges the difference between scholarship that

incorporates the impact of animal life on humans but is essentially focused
on humans, and that which aims ‘to explore the history of nonhuman
animals as subjects in their own right and for their own sakes’.2 Nearly all
medical history scholarship falls into the former category. While animals
do feature in it, and to an increasing degree since the turn of the twenty-
first century,3 they are usually shadowy, marginal creatures, ‘mere blank
1 Kirk
and Worboys (2011), p. 561.
2 Benson(2011) p. 5.
3 Recent reviews of this literature include: Rader (2007), Kirk and Worboys (2011),
Woods (2016, 2017a, 2017b), Kirk (2017).
© The Author(s) 2018 1

A. Woods et al., Animals and the Shaping of Modern Medicine,
Medicine and Biomedical Sciences in Modern History,
https://doi.org/10.1007/978-3-319-64337-3_1
2 A. Woods et al.
pages onto which humans wrote meaning’.4 They appear because of their
implications for human health and medicine, or because of their capacity
to illuminate wider developments in human history, such as the growth of
government, colonialism and international trade.
This volume breaks new ground in applying Benson’s second per-
spective to medical history: ‘to explore the history of nonhuman ani-
mals as subjects in their own right and for their own sakes’.5 Humans
remain important, of course, for ultimately we can only know about
animals from the records that humans have created, and which reflect
human interests in animals. However, by taking animals seriously as his-
torical subjects, it is possible to shed a different light on human history
by revealing the myriad ways in which animals have influenced human
actions and perceptions. Adopting this approach also illuminates ani-
mals as creatures with their own histories, which have been profoundly
altered by their relationships with humans, and the roles that humans
have decided they should perform. It results in a richer, less anthropo-
centric account of the medical past, which reveals how, in different times
and places, animals have experienced medicine, how they have been pro-
duced by it and how they have changed it.
In widening the historical lens to incorporate animals and their fash-
ioning into medical subjects and objects, this volume pursues three key
goals. First, it seeks to make a programmatic contribution to the field of
medical history by elucidating some of the largely unrecognized ways in
which animals have informed the knowledges, practices and social forma-
tions of medicine. Through analysing key contexts in which animals have
attracted medical attention and with what effects, it will expose a series of
medical problems, concerns, personnel and practices that barely feature
in existing scholarship. In addition, by studying the historical position-
ing of animals at the shifting boundaries between medicine, veterinary
medicine and the life sciences, it will cast new light on the relationships
between these fields. It will thereby demonstrate how, by attending to
the more-than-human dimensions of medicine, we reach new under-
standings of its historical identity, participants and manner of pursuit.
Second, the volume seeks to enhance the burgeoning field of animal
history by offering the first substantive account of animals in medicine.
4 Fudge (2006).
5 Benson (2011) p. 5.
1 INTRODUCTION: CENTRING ANIMALS WITHIN MEDICAL HISTORY 3
Expanding beyond much-studied laboratory contexts to explore the

medical history of animals in zoos, on farms, in hospitals, post-mortem
rooms and international policy arenas, it illuminates the diverse species
that have participated in medicine, the many roles they have played in
it, and how their bodies and habits have both shaped and been shaped
by its ideas, practices and institutions. Crucially, the volume highlights
how these diverse species forged multispecies networks, thereby extend-
ing animal history’s typical focus on the dyadic relationships between
humans and another species of animal.
The third objective of this volume is to speak to the twenty-first-cen-
tury initiative known as One Health (OH). Featuring prominently in
medical, veterinary and scientific publications, and in national and inter-
national health policy and position statements, OH pursues an expansive
vision of improving health and wellbeing through the multidisciplinary
study of problems at the interface of humans, animals and their environ-
ments. For its proponents, OH represents a necessary response to a host
of shared threats to human and animal health, such as emerging diseases
that transmit between animals and humans, antimicrobial resistance,
food insecurity, food safety and climate change. They argue that such
issues cannot be tackled effectively within the traditional disciplinary
compartments of human medicine, veterinary medicine and the life sci-
ences. Rather, integrated, coordinated approaches are required, in which
the health of animals is considered in relation to the health of humans
and the environment.6 This volume situates OH within a longer histori-
cal context by illuminating certain precedents to this way of working. It
also offers a critical, empirically grounded perspective on its operation
today by exploring the circumstances that gave rise to its emergence as
a self-conscious movement, and how its proponents conceptualize the
roles of animals within it.
In addressing these three objectives, the volume also addresses three
distinct audiences: historians of animals, historians of science and medi-
cine, and health professionals concerned with OH today. The remainder
of this chapter introduces the history of animals as a field of enquiry, and
situates this study in relation to it. While historians of animals will be
familiar with its discussion of the methodological and conceptual issues
6 For example: Gibbs (2014), American Veterinary Association, Centers for Disease
Control and Prevention, One Health Initiative.

4 A. Woods et al.
that are important to the writing of animal histories, we present them

here for the benefit of medical historians who may not be familiar with
them. We also provide an important overview of how we approach the
history of animals in medicine, and conclude by elaborating on the vol-
ume’s objectives, outlining its research questions and introducing the
case studies that follow.
1.1 Why Animals?

The ways in which non-human animals have shaped human history is a
pressing and important issue for historians today. Recent years have wit-
nessed increasing engagement with the subject, manifesting in a bur-
geoning body of literature that draws on perspectives from material
culture studies, science and technology studies, zoology, performance
studies, and environmental, social and cultural history. Directing their
attention to a variety of animal species, authors have addressed the lives
and experiences of animals, their categorization and manipulation by
humans, their relationships with humans and environments, and their
representations within art and literature.7 The eclectic methods of animal
history, and the many differences between its animal subjects (some of
which had more historical similarities with humans than with each other)
have led some historians to ask whether it can be said to constitute a
coherent field of enquiry, or whether it primarily offers an approach to
animals which can be applied to all existing types of history.8 Insofar
as animal history is a field, this volume is intended to be a contribu-
tion to that body of work, but it also draws on animal history ideas and
approaches in order to develop new perspectives on medical history.
Animal historians acknowledge that the significance of animals to war,
agriculture, science, colonialism, sport and the environment means that
they have long featured in scholarly histories, but usually as supporting
7 Seminal works include: Ritvo (1987), Kete (1994), Anderson (2004). Valuable edited
collections include: Henninger-Voss (2002), Rothfels (2002), Kalof and Resl (2007), Brantz
(2010), Shaw (2013a, 2013b), Nance (2015). An even larger scholarship addresses the contem-
porary dimensions of human–animal relationships, drawing on disciplines such as philosophy,
anthropology, geography, English literature and cultural studies. For an introductory overview,
see Marvin and McHugh (2014). Key authors who have set the framework for thinking about
these issues include: Agamben (2002), Derrida (2002), Wolfe (2003), Haraway (2008).
8 Swart (2007), Andrews (n.d.).
actors in the drama of human history. Their stated intention is to bring

animals in from the margins and position them at the centre of histor-
ical analysis in order to explore the intertwining of human and animal
lives, and the development of human ideas about, and relationships with,
animals.9 Conceptualizing animals as creatures with their own histories
and the unintentional capacity to effect historical change, authors seek to
trace ‘the many ways in which humans construct and are constructed by
animals in the past’.10 The purpose of these analyses is not simply to fill
a gap in the writing of human history but to rethink conventional histo-
riographies. This volume follows their lead in acknowledging animals as
shapers of medicine in history, and also as shapers of the ways in which
we, as scholars, perceive and write about medical history.11
Developments beyond the academy have helped to precipitate this
‘animal turn’. Since the later twentieth-century, animal-related causes,
from opposition to factory farming and animal experimentation, to the
wider improvement of animal welfare, have gained increasing public
support and political traction. Portrayed variously as victims of, or con-
tributors to, environmental degradation, animals have also become a
key aspect of wider concerns about human interactions with the natu-
ral world. Meanwhile, the impacts of diseases such as bovine spongiform
encephalopathy (BSE) and avian influenza, which emerged in ani-
mals and spread to people, have made the health connections between
humans and animals more visible and threatening. These developments
have prompted much reflection on human responsibilities for non-
human others, and how to live sustainably with them.12 They have also
encouraged animal scientists such as ethologists, vets and ecologists, to
study the sentience and subjectivities of animals, and their relationships
with their environments.13
Enhanced concern for animals and human–animal relations in the pre-
sent has helped to draw attention to their pasts within different domains
of historical scholarship. Situating animals within nature as constituents
9 Ritvo (2002), Fudge (2002), Kean (2012), Sivasundaram (2015).

10 Fudge (2006).
11 We thank Tamar Novick of the Max Planck Institute for the History of Science, Berlin,
for making this observation.

12 Ritvo (2002), Shaw (2013a, 2013b), Vandersommers (2016).
13 For example: Bekoff (2002), Grandin and Johnson (2005).

6 A. Woods et al.
of environments and ecosystems, environmental historians have explored

the interplay between ‘nature’ and ‘culture’ in the shaping of human
pasts.14 Challenging the very notion of a nature–culture dichotomy
and the priority it awards to human ‘culture’ over animal ‘nature’, post-
humanist scholars have sought to understand how human and animal
differences, essences and linkages have been constructed through his-
torically specific encounters.15 By contrast, for social and cultural histo-
rians, the fact that ‘humans are always, and have always been, enmeshed
in social relations with animals’16 calls for a social historical approach to
animal histories. Their work has established animals as the latest ben-
eficiaries of ‘history from below’, a genre that originated in the 1970s
with E.P. Thompson’s history of the working classes, and expanded
subsequently to incorporate other neglected historical subjects such as
women, colonized peoples, marginalized ethnic groups and the mentally
ill.17 The animals studied tend to be celebrity animals, charismatic wild-
life and those domesticated species that have entered into close relation-
ships with humans. Other, more marginal creatures have been neglected.
This volume offers a partial corrective by examining the medical histories
of some uncharismatic and historically overlooked animals, such as tape-
worms and farmed livestock.
As with the other groups targeted by ‘history from below’, there is
an explicit political dimension to much animal history writing. Some
scholars, who locate themselves within the field of critical animal stud-
ies, aim to improve animal lives in the present by uncovering and criti-
cizing the ways in which humans have exploited them in the past.18
For other animal historians, these narratives of animal domination and
oppression are too simplistic. They, too, are often keen to effect pre-
sent-day changes in attitudes to animals by revealing their treatment in
the past. Consequently, they remain alert to the power dynamics that
have informed past human–animal relationships.19 However, they also
emphasize the complexity and historical specificity of those relationships,
14 McNeill (2000), Nash (2005), Cronon (1990).

15 Lorimer (2009), Cole (2011).
16 Philo and Wilbert (2000) p. 2. See also Swart (2007) p. 276, Eitler (2014) p. 262.
17 Ritvo (2002), Kean (2012). For a history of animals as workers, see Hribal (2007).
18 Taylor and Twine (2014), Institute for Critical Animal Studies.
19 Fudge (2012).
and the ‘resistance, compliance and compromise’ that variously charac-

terized them.20 This volume follows their lead in believing that historians
have a moral obligation to document how human actions have impacted
animal lives. It seeks to develop a nuanced understanding of the human–
animal relationships that were forged through medical research and prac-
tice, and to analyse the implications of those relationships for humans,
animals and medicine.
1.2 Writing Animal Histories

There are particular challenges associated with writing the history of ani-
mals. Some scholars have questioned whether it is even possible. Pearson
and Weismantel state that the challenges are at once ontological (‘a ques-
tion of imagining animal being’), epistemological (because our commu-
nications with animals are non-verbal) and methodological (how can we
write authentic animal histories when the only records that survive of
them were created by humans?). They suggest that these difficulties are
linked: if animals have no voices, they cannot leave records from which
historians can write their pasts; without language, consciousness or inten-
tionality they are unable to participate in history except as subjects of
biological evolution. For some commentators, it follows that the study of
animals belongs within the natural sciences not the humanities—which
are by definition concerned with human society and culture.21 This
stance has roots in Christian theology, which asserts human dominion
over animals, and in Enlightenment thinking, which posits fundamental
distinctions between humans and animals, nature and culture.22
These dichotomies and the exceptional status they award to humans
have been challenged by post-modern and post-humanist scholarship,
and by studies of developments within technoscience, food production
and climate science, which provide concrete examples of the impossibil-
ity of separating nature from society.23 Historical analyses lend support
to these challenges. By illuminating circumstances in which particular
20 Pooley-Ebert (2015) p. 165.

21 Pearson and Weismantel (2010).
22 Thomas (1983).
23 Latour (1993), Mitchell (2002), Schrepfer and Scranton (2003), Agamben (2002),
Lorimer (2009).
8 A. Woods et al.
humans (notably women, the mentally ill and certain ethnic groups)
were invested with animal characteristics, and certain animals (notably
horses, dogs and primates) were awarded privileged, quasi-human sta-
tus, it reveals that human–animal boundaries are not fixed in nature but
shaped by human society and culture. Therefore animals and animality
are not distinct from, but constitutive of the human, in ways that change
over time and place.24
If the human–animal boundary is blurred, then where does this leave
assertions that without cognition or conscious thought, animals cannot
be active historical subjects? Scholars have tackled this question by con-
sidering whether the ability to act on history can exist separately from
the intention to do so, and whether animals can be historical subjects
without necessarily possessing subjectivity. They suggest that while ani-
mals do not understand or care about human knowledge, politics and
cultures, nevertheless they can make a difference to them. Therefore, in
principle, animals do possess agency—where agency constitutes the abil-
ity to act.25 For animal historians, the nature of that agency is an empiri-
cal question. It is not a natural, permanent attribute; rather, it emerges
and is performed through social relationships, which vary by time, space
and species.26
In locating animal agency within human–non-human encounters,
many scholars draw on ideas associated with Actor Network Theory
(ANT). Developed within the field of science studies by Bruno Latour,
Michel Callon and John Law in the 1980s, ANT sought to remove dis-
tinctions between nature and society. It represented a critical response
to prevailing understandings of scientific knowledge and technical inno-
vation as either realist or sociological.27 It has since been adopted and
developed by scholars in many other disciplines. ANT has many variants
but it essentially proposes a relational way of understanding the world. In
widening the traditional analytical lens to incorporate interacting human
and non-human entities, it opens up a space for animals as historical
actors. ANT presents agency, and actors, as products of the unstable web
24 Ritvo (1995), Fudge (2002), Ritvo (2007), Hochadel (2010), Bourke (2011), Davis
(2014).
25 Fudge (2006), Law and Mol (2008), McFarland and Hediger (2009).
26 Philo and Wilbert (2000), Eitler (2014), Pooley-Ebert (2015).
27 Callon and Law (1986), Latour (1988, 2005).

of relations (or networks) in which all are embedded. The configuration

of these networks affects how the capacity to act is distributed among
and deployed by its members. Their actions reshape ideas and practices,
bodies and materials, experiences and social relations, and, by extension,
the network itself. In this formulation, it is not only the animal’s capacity
to effect historical change but also the animal itself which is constantly
being reconfigured through its embeddedness in multiple interactive,
ever-changing relationships.28
Animal studies scholars frequently cite ANT when reflecting on the
agency of animals. However, their recognition that pre-existing power
differentials, social contexts and species differences have an important
bearing on network formation, the relationships of its participants, and
consequently on the animal’s ability to act, seems to support criticisms
of ANT’s ‘flat ontology’—its refusal to acknowledge any differences
between actors other than those created through networks.29 While we
agree with such criticisms, we, too, find ANT to be a useful heuristic
device because of the significance it awards to relationships. It suggests
that through scrutinizing ‘the entire lived experience of quotidian and
extraordinary interactions—embodied and imaginary, material and sym-
bolic—that occur within space and in particular locations, and involve
humans and animals in multiple forms of engagement and exchange’,30
we can work out how animals have changed medicine and have, in turn,
been changed by it.31 Moreover, by privileging relationships, as ANT
exhorts us to, we are able to identify the interconnections between mul-
tiple species.32
Thinking about animals in history has also been profoundly influenced
by Donna Haraway’s writing. While acknowledging the uneven distribu-
tion of pain and suffering between human and animals, she perceives
their relationships to be characterized by mutual adaptation rather than
exploitation, and seeks to determine how, in historically and culturally
specific circumstances, humans and animals come together in ‘material-
semiotic nodes or knots in which diverse bodies and meanings coshape
28 Pearson (2013) pp. 128–45, Eitler (2014).

29 Barron (2003), McLean and Hassard (2004).
30 Pearson and Weismantel (2010).
31 Kean (2012).
32 Pettit et al. (2015).

10 A. Woods et al.
one another’.33 As Law and Mol point out, this does not mean that ani-
mals have the capacity to control history, just that by entering into rela-
tionships with human and other non-human actors they are able to make
a difference to it. Their analysis of Cumbrian sheep during the British
2001 foot-and-mouth disease epidemic offers some concrete examples.
The variable bodily responses of sheep to encounters with the foot-and-
mouth disease virus affected whether the vets that inspected them were
able to detect and respond to it. The hill sheep’s ability to know its place
or ‘heft’ on the hills fashioned the landscape in ways that were valued by
humans, and which led ultimately to their exemption from the govern-
ment’s contiguous culling policy. These outcomes were not necessarily
predictable: they resulted from the unstable, indeterminate webs of prac-
tices that drew together and were created by sheep and their co-actors.
Law and Mol conclude that the important question is not whether but
how animals act.34
We will address this question by analysing the roles that animals have
performed within medical research and practice. Some of these roles will
be familiar to medical historians. In laboratories, animals were fashioned
into experimental subjects and manipulated to cast light on health and
disease. Animals that fell sick performed roles as disease victims, and
were sometimes transformed into patients. Alternatively, or addition-
ally, they were regarded as hosts and potential transmitters of infection.
Animals have also performed other, less well-documented roles in medi-
cine: as pathological specimens, points of comparison with other species,
commercial products, shapers and victims of food systems and natural
environments, and vehicles for personal and professional advancement.
They have often performed several roles simultaneously or sequen-
tially throughout their lives and afterlives. Roles have reshaped animals,
physically and conceptually, and therefore impacted on their histo-

ries. Roles also had their own histories that were influenced partly by
human–animal relationships and their disruption by disease, and also by
prevailing ideas of disease and the tools available to conceptualize, inves-
tigate and manage it. While it was humans who awarded roles to ani-
mals, animals were not passive recipients. Their bodies, behaviours and
relationships with humans have invited the awarding of particular roles
33 Haraway (2008) p. 4.
34 Law and Mol (2008) p. 74.
and influenced their performance within them. Different roles have pro-
vided animals with different opportunities to shape medicine, with rami-
fications for the health of both humans and animals. The concept of the
animal role therefore offers a useful tool for illuminating the historical
co-constitution of humans, animals and medicine.
1.3 Animals in Medical History

Inspired by Haraway’s thinking, in 2011, Kirk and Worboys called for
the history of medicine to be rewritten as a history of interspecies inter-
actions.35 However, to date, few scholars have adopted this approach.
There are only a handful of key works that foreground the relationships
between humans and animals, and these focus narrowly on experimental
settings.36 Studies of the co-constitution of animals and medicine are also
few in number and derive more from the history of the life sciences than
medical history.37 For the most part, animals fade into the background
of medical history writing.38 Scholars influenced by the ‘animal turn’
have not been drawn to study the history of medicine, while historians of
medicine have remained largely unaware of the ‘animal turn’.
This situation has not arisen because animals were unimportant to
medicine. On the contrary, as this volume and its extended bibliogra-
phy demonstrate, they have been integral to its history. Medical scien-
tists employed animals to develop new knowledge of bodies, minds and
diseases; to generate biological products, and to test the safety and effi-
cacy of drugs. Animals supplied nutrition to humans and transmitted
diseases to them. The state of animal health powerfully influenced—and
was influenced by—their environments. Animals were treated as patients,
fashioned into pathological specimens, and their diseases compared across
species. So far, however, these animal roles have been studied from a
largely human perspective. Their capacity to shape medicine has attracted
little attention, and they have been rarely studied as medical subjects ‘in
their own right and for their own sakes’—not even within the field of
35 Kirk and Worboys (2011).

36 Rupke (1990), Todes (1997), Dror (1999), Guerrini (2003), Schlich and Schlünder
(2009), Kirk (2014).
37 Clarke and Fujimura (1992), Clause (1993), Kohler (1994), Rader (2007).
38 Woods (2017b).
12 A. Woods et al.
veterinary history, which is ostensibly focused on the health of animals as

a problem in its own right.39
We can offer only tentative explanations for why animal histories of
medicine have yet to be written. It may be because of the dominance of
cultural studies’ perspectives within animal and human–animal history,
which have tended to foreground animals’ cultural and symbolic roles in
society rather than delving into the content and material practices of ani-
mal science and medicine. It may also result from a preference for writing
the histories of individual animals that are visible in today’s society and
with which humans develop empathetic bonds: pet animals, zoo animals
and charismatic wildlife species. Farm animals, which have wielded con-
siderable influence over human health, are comparatively neglected. Such
explanations do not apply to historians of medicine. The field’s long-
lasting engagement with ANT,40 its preoccupation with the histories of
bodies and material practices, the scientific training of many of its schol-
ars, and their expertise—generated through writing patient histories—in
thinking about medical history ‘from below’, means that the tools for
writing animal-centred histories of medicine are already in circulation.41
The failure to apply them may be due to the anthropocentrism that char-
acterized most fields of history writing until relatively recently. However,
we believe that disciplinary traditions may also be to blame.
It is now more than 20 years since the late Roy Porter asserted that,
‘in the academic world, it is automatically assumed that a “historian of
medicine” is a person who works on the history of human medicine’.42
He attributed this assumption to modernist notions of human differ-
ence from, and primacy over, animals. While in the intervening years
these notions have come increasingly under attack, scholars have not sig-
nificantly revised their perceptions of what constitutes medical history.
There remains an implicit assumption that human health and medicine
lie at the heart of this field. Sick animals whose health had no direct con-
sequence for humans are relegated to the small subfield of veterinary
39 Benson (2011) p. 5. Susan Jones is one of the few veterinary historians to foreground
animals and their relationships with humans. A good example is Jones (1997). See also
Degeling (2009).
40 Latour (1988).
41 Porter (1985), Beier (1987), Warner (1999), Gillis (2006), Cooter (2010), Crozier
(2010), Hurren (2012).
42 Porter (1993) p. 19.
history, while other aspects of animal life are studied by historians of

biology. We hold that this compartmentalization is artificial and unhelp-
ful. It does not reflect the historical positioning of animals within these
fields, and it produces a narrowly anthropocentric framing of ‘medicine’
which is frequently at odds with its historical identity.
This volume challenges such conceptions by revealing that mod-
ern medicine, as developed in the West over the last two centuries,
was a more-than-human endeavour, whose boundaries with veteri-
nary medicine and biology were porous and in a constant state of flux.
In certain historical contexts, animals contributed to the compartmen-
talization of these domains. In others, they helped to break down the
barriers between them, particularly through their investigation within
boundary-crossing fields such as parasitology, zoology, comparative
medicine, nutrition and agriculture.43 Animals contributed to the forma-
tion of these fields, and were, in turn, formed by them. By elucidating
these processes, this volume not only sheds light on the history of ani-
mals. In identifying the ideas, methods, problems, places and people who
engaged with their health, it also develops a new perspective on medicine
itself—and therefore on what constitutes the field of medical history.
In developing this animal-centred medical history, we have had to
engage with the tricky issue of how to write the histories of non-ver-
bal creatures when the only records that survive of them were created
by humans. Some scholars have attempted to overcome this problem
by using modern scientific understandings of animals to retrospec-
tively interpret their behaviours and experiences.44 As historians of sci-
ence and medicine, we find this approach deeply problematic, because
in granting a timeless universality to scientific interpretations that are in
fact products of specific historical circumstances, it reifies the nature–
culture divide.45 Other scholars have argued that the problem cannot
be overcome; that it is impossible to truly ‘know’ the authentic histori-
cal animal because surviving records are mere cultural representations
of them.46 Swart has challenged this view. Questioning the very notion
of authenticity, she points out that the ‘facts’ about animals are always
43 Woods (2017c).
44 For example: Pearson (2013), Foote and Gunnels (2015), Pooley-Ebert (2015).
45 Latour (1993).
46 Fudge (2002) p. 6.
14 A. Woods et al.
human interpretations, not ‘real’ accounts of them.47 Benson goes fur-

ther by arguing that because animals have informed the human produc-
tion of records of them, such records—or traces—are more than cultural
representations; they comprise ‘material-semiotic remnants’ of animals, a
nexus of language and materiality that offer ‘unintentional indexes of a
now-absent presence’.48
Inspired by Benson’s thinking around animal traces, we approach
medical historical records of animals as evidence of their interactions with
humans, whose analysis reveals how animals made a difference to, and
were changed by, medicine. We perceive multiple layers of animal traces
which gained meaning in reference to each other. There are the imme-
diate material remains of diseased animal bodies such as taxidermy and
museum specimens. Then there are the narratives, statistics and images
that humans have created from these materials and from other long-dead
animals whose remains were not preserved. Finally, there are the knowl-
edge-practices and social relationships that were fashioned by and from
these creations. In studying these traces, we ask not only what they reveal
about the health of animals in history and the roles that animals played
in medicine. By reflecting on why this ‘animal archive’ was created in
the first place, we also learn about the animal’s capacity to attract human
attention, and the relationships that bound them.49
1.4 One Health and its Histories

One of the inspirations and intended audiences for this work is the con-
temporary movement for OH, whose features and early twenty-first-cen-
tury emergence are explored in Chapter 6. OH is underpinned by the
belief that some of the most important health threats faced today are not
species specific, and consequently can only be tackled by interdisciplinary
working across the domains of human medicine, veterinary medicine and
the life sciences. Its integrated approach to human and animal health
implies that medicine cannot achieve its goals through a purely anthro-
pocentric approach. Just as other contemporary agendas have prompted
historians to study animal pasts, so has the push for OH drawn our
attention to the roles played by animals in the history of medicine.
47 Swart (2015).
48 Benson (2011) p. 3.
49 Tortorici (2015).
While the term OH is new, the concept is not. This is well recog-
nized by OH advocates, who use historical observations of human–ani-
mal health connections in attempts to legitimize their agenda, and win
momentum and funding for it.50 They locate the roots of OH in an
earlier agenda known as One Medicine (OM), which developed in the
1960s and 1970s with the aim of bringing human and veterinary medi-
cine into closer alignment.51 OM was, in turn, presaged by work dat-
ing back to the nineteenth century in the fields of comparative medicine
and veterinary public health, where human and animal health prob-
lems were considered in tandem.52 OH advocates offer a single, highly
selective interpretation of this history. They highlight the work of key
high-profile figures whose work crossed the human–animal divide and
had a lasting impact on medicine: Louis Pasteur, Robert Koch, William
Osler, Theobald Smith, John McFadyean and Rudolph Virchow (who
is frequently quoted, in the absence of any identifiable source material,
as stating that ‘between human and animal medicine there is no divid-
ing line’).53 Extracting these individuals from their historical contexts,
OH advocates present them as far-sighted geniuses who recognized the
merits of a OH approach. There is little understanding of what, within
the context of the time, these men thought they were doing and why;
whether their work was typical or unusual; and how animals might have
shaped, or been shaped, by it.
One of the goals of this volume is to develop a more critical, evidence-
based, animal-centred history of OH by documenting and explaining
the circumstances in which animals and their diseases became important
to human medical agendas. By identifying what motivated their investi-
gation by a range of lesser-known individuals, how these investigations
were pursued in policy and practice, and with what implications for
human and animal health, the book will illuminate precedents to OH
today, and draw insights of relevance to its future operation. It will also
offer an explanation for the emergence of OH as a self-conscious twenty-
first-century agenda and unpack the place of animals within it.
50 Woods and Bresalier (2014).

51 Schwabe (1984), Zinsstag et al. (2011).
52 Bresalier et al. (2015).
53 Schwabe (1984), Michell (2000) pp. 101–6, Day (2008) pp. 151–3.
16 A. Woods et al.
We do not attempt a complete account of the history of animals in

medicine: the subject is too vast and too neglected.54 Instead, we aim to
set a new research agenda and to illustrate where it might lead through a
series of case studies that are drawn from the authors’ individual research
programmes. These studies explore animal histories of medicine, and
histories of animals in medicine, on scales ranging from the local to
the global, from the 1830s to the present day. They are designed to be
read separately as standalone examples of the contexts in which animals
became important to medicine and the difference they made to it. Read
collectively, they illuminate the diverse species, spaces, methods, peo-
ple, problems and contexts of enquiry that were involved in construct-
ing animal health as a medical problem. In tracing how that construction
changed over time, they also trace the shifting place of animals within
medicine, and how animals moulded its relationships with veterinary
medicine and the life sciences.
Cross-cutting questions include: What circumstances attracted human
attention to animals and their diseases, and what networks developed
around them? How can we account for the attention paid to animals by
members of the medical profession, which is generally assumed to be
exclusively concerned with the health of humans? How and where were
investigations and interventions performed on animals, and what roles
did animals play within them? While our human protagonists’ affec-
tive relationships with animals are generally impossible to discern, the
concept of the animal role permits analysis of the multiple values that
humans assigned to animals. We also ask: What difference did animals
make to medicine—to its ideas, practices, the health of its human and
animal subjects, and its interpersonal and interdisciplinary relationships?
What difference did medicine make to animals—to their bodies and
experiences in life, the manner and timing of their deaths, and to their
afterlives?
In selecting which aspects of the history of animals in medicine to
study, we made a deliberate decision not to focus on the laboratory-
based subjects of experimental medicine. This is partly because these
animals, notably rodents and dogs, already feature in the handful of
existing animal-centred accounts of medical history. It is also because
54 For a chronological overview, see Bresalier et al. (2015).

experimental medicine is already overrepresented within medical history

scholarship. The widely held perception that the pursuit of knowledge
through experiments is one of the defining features of modern medicine,
means that scholars have dedicated considerable attention to its history,
starting with the growth of experimental physiology around 1800, and
progressing to the emergence of other experimental sciences: bacteriol-
ogy in the 1860s and 1870s, pharmacology, endocrinology, nutrition sci-
ence and immunology in the decades around 1900, and biomedicine in the
post-Second World War era.55 The attention lavished on the topic is such
that many scholars seem to assume by default that the history of animals in
medicine is a history of experimental animals, or ‘animal models’ of disease,
that were manipulated in laboratories for the benefit of human health.
This volume seeks to dislodge that perception through a series of case
studies that decentre not only humans but also laboratory-based experi-
mentation from the history of modern medicine. The studies address
a series of other important medical contexts whose histories have been
obscured by the historiographical focus on the laboratory: natural history,
zoology, parasitology, comparative anatomy, ecology, nutrition and agri-
culture. Lying at the borderlands of human medicine, veterinary medicine
and the life sciences, and shedding light on their shifting points of intersec-
tion, analysis of these contexts shifts the historical focus away from labo-
ratory rodents and dogs towards a wide array of other species that hardly
ever feature in accounts of medical history: zoo animals, Scottish sheep,
cows of the developing world, and the tapeworm, Echinococcus granu-
losus. These animals performed various roles, including, but not confined
to, those of disease victims, patients, experimental material, shapers and
products of their environments, hosts and transmitters of infection, tools
for thinking comparatively across species, spontaneous (as opposed to
experimentally designed) models of disease, and sources of human nutri-
tion. Their investigation relied not only on experimental practices but
also on observation, categorization, comparison, statistical analysis and
clinical trials. These methods were applied far beyond the laboratory, in
animal houses, post-mortem rooms, hospitals, farms, field stations and
slaughterhouses. By drawing our attention to these historically neglected
aspects of medicine, the animals studied in this volume also expand and
55 Guerrini (2003), Lowy (2003), Rader (2007), Schlich et al. (2009), Löwy (2011),
Franco (2013).
18 A. Woods et al.
alter received conceptions of what constituted medicine in history, and who

were its key actors.
The first case study, presented in Chapter 2, is situated in Britain’s
mid-nineteenth-century zoological gardens, particularly those located
in London and Dublin. It documents the diseases and deaths that beset
their diverse animal inhabitants, and argues that as a consequence, the
zoos became important sites of medical research and practice. It shows
how medical men who helped to run the zoos, and medical visitors who
hoped to make their names within them, used their knowledge and prac-
tice of human medicine and comparative anatomy to advance the health
of zoo animals, and devise comparative pathological understandings of
their diseases. These men awarded animals the roles of patients, victims
of their environments, pathological specimens and points of interspecies
comparison. They manipulated animal bodies, surroundings and man-
agement in ways that were shaped by animal biologies and behaviours.
Through these activities, the zoo became medical, and medicine zoologi-
cal. An array of vertebrate species fell under the medical gaze, and helped
to generate new knowledge of health and disease that found applications
in human medicine.
In Chapter 3 the focus shifts to diseased and dying sheep on farms
in and around Scotland at the turn of the twentieth century. It reveals
how these animals came to be regarded as victims of their environ-
ment, and positioned at the hub of a research network containing
farmers, doctors, vets, natural historians and zoologists. It examines
the investigations performed by this network, and how sheep fash-
ioned and were fashioned by them. It then describes and explains key
changes to the network, which shifted the location of investigations
from farms to laboratories, and distanced doctors and practical farm-
ers from the scientific study of sheep. Awarded new roles as hosts and
transmitters of infection, sheep lost influence over investigators’ activi-
ties. Meanwhile, veterinarians sought to capture sick sheep for them-
selves by claiming superior knowledge that derived from their unique
relationships with them. In these ways, sheep first integrated, and then
contributed to widening divisions between the various experts in their
diseases.
Chapter 4 is concerned with diseased and undernourished dairy cat-
tle, and how they came to be perceived not simply as threats to farm-
ing profits but as contributors to world hunger and ill health. Moving
from interwar Britain and its empire, to the post-war international

stage, it explores how developments in nutritional science and veterinary
medicine, combined with economic depression, food shortages and the
effects of war, drew attention to the undernourished, unhealthy bodies
of both humans and cows, and suggested connections between them. By
the early 1950s, under the United Nations and its agencies, cows had
become key participants in the campaign against human hunger in the
developing world. Their unproductive bodies inspired the formation of
new health structures that brought together experts in human health,
nutrition, veterinary medicine and agricultural science to create new
types of cow that would prove more capable of supporting human health
and nutrition.
Chapter 5 continues to cross borders between nations and disciplines in
its study of the tapeworm, Echinococcus granulosus, and the ideas and inves-
tigations it inspired. It is particularly concerned with the work of parasi-
tologist Calvin Schwabe, who is better known as a progenitor of the recent
movement for OH. For Schwabe, Echinococcus was an animal in its own
right, an active, opportunistic participant in both human and non-human
ecological and cultural interactions. In following Schwabe as he followed
Echinococcus, from the laboratory into human communities and multi-
species ecosystems, and from Beirut to Kenya to California, this chapter
reveals that the roots of his commitment to unifying human and veterinary
medicine lay in his deep-seated engagement with the parasite. His investi-
gations into its body and behaviour led him to view distinctions between
human and non-human species as culturally contingent rather than fun-
damentally biological. This fed his conviction that human and veterinary
medicine could only function truly effectively when practised in tandem.
Chapter 6 takes forward the story of OH by exploring its emer-
gence as a self-conscious movement, dedicated to the integrated study
of problems at the interface between human health, animal health and
the environment. It explores how Schwabe’s work influenced, and was
reconfigured by, this movement, and locates its early development in
several research and policy networks, which produced not one but sev-
eral different forms of OH. The chapter also examines how human–
animal health relationships have inspired and shaped OH, and how they
are represented—in sometimes contradictory ways—in the texts and
images produced by its researchers and advocates. It argues that in priv-
ileging the roles of animals as transmitters of diseases to humans, and
20 A. Woods et al.
experimental models of human diseases, OH rebrands longstanding

research agendas that are far more concerned with the health of humans
than that of animals.
Chapter 7 concludes by summarizing what these chapters have revealed
about the medical history of animals and the animal history of medicine.
It reflects on the implications of these findings for how historians think
about and study the history of medicine, and for how OH advocates con-
ceptualize and pursue their integrating agenda. The volume ends with
an annotated bibliography of animals in the history of medicine, which
offers an entry point for scholars who are new to the field and is organized
around some of the key animal roles that are explored in the chapters.
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CHAPTER 2
Doctors in the Zoo: Connecting Human

and Animal Health in British Zoological
Gardens, c.1828–1890
Abigail Woods
In 1865, the Royal Zoological Society of Ireland (RZSI) announced
the death of a three-year-old male Indian rhino in its Dublin Zoological
Gardens. The event was a considerable blow to the society, which had
paid £160 for the unusual creature in the hope of inspiring scientific
and public interest in the zoo. However, on arrival from Calcutta just
eight months previously, the rhino was already sickly and suffering from
fits. It was attended by three medical members of the RZSI, including
the secretary, Reverend Professor Samuel Haughton, of Trinity College
Dublin (TCD), who recommended the administration of three pints of
boiled rice with bran, and a gallon of milk with some tonic mixed in.
However, the fits continued. Haughton elected to increase the dose of
tonic and remove cabbage from the diet. This brought about a tempo-
rary improvement, but in April 1865 the animal was found in pain with
a prolapsed rectum. Haughton, two other doctors and two vets were
summoned. They administered castor oil, opium, aromatic spirits of
ammonia and turpentine, but to no avail.1
After its death the rhino continued to attract attention. The Royal
Dublin Society offered £15 for the body in the hope of adding to its col-
lection of comparative anatomy specimens. It was outbid by Haughton,
who habitually dissected animals that died in the zoo.2 His post-mortem
1 de Courcy (2010). See also de Courcy (2009).

2 de Courcy (2010), Proceedings of the Royal Zoological Society of Ireland (1863–1864) p. 13.
© The Author(s) 2018 27

A. Woods et al., Animals and the Shaping of Modern Medicine,
Medicine and Biomedical Sciences in Modern History,
https://doi.org/10.1007/978-3-319-64337-3_2
28 A. Woods
examination of the rhino’s pathology and anatomy was attended by

anatomists, medical men and the queen’s veterinary surgeon for Ireland,
with assistance provided by medical students and the demonstrator in
anatomy at the Royal College of Surgeons, Ireland. By then, the rhino
had begun to decompose and the stench from its body was almost intol-
erable, causing several of the attendants to suffer typhoid diarrhoea.
Examination revealed that its rectum had prolapsed and ruptured, and its
stomach was distended almost to bursting with fermenting Indian corn.
This had exerted pressure on the diaphragm, leading to death by suffoca-
tion. A furious Haughton instructed the council of the RZSI to institute
a searching enquiry as to why the rhino had been fed corn when it was
not listed on the society’s formally prescribed dietary.3 Proceeding to dis-
sect the remainder of the body, he wrote a lengthy report on the rhino’s
muscles, which he published alongside his pathological findings in the
Proceedings of the Irish Academy. As number 16 in his series of 18 ‘Notes
on Animal Mechanics’, the report informed his 1873 volume, Principles
of Animal Mechanics, which compared and contrasted the bodies of vari-
ous species including humans, and argued—contrary to Darwin—for a
teleological view of nature.4 What remained of the rhino was sent to a
taxidermist and then displayed alongside other animals in the zoological
museum of TCD.5
This vignette of the life, death and afterlife of the unfortunate Dublin
rhino introduces several key themes that will be investigated further in
this chapter. First, it offers a glimpse of the illness experiences of animals
that were confined to nineteenth-century zoological gardens for their
frequently short and sickly lives. Second, it reveals the sorts of human
responses that those illnesses inspired. Sick animals were fashioned into
patients, pathological specimens, victims of their environments and points
of interspecies comparison, and subjected to medication and dietary
modifications in life, and dissection after death. Third, it shows that these
responses were led not, as one might expect, by veterinary surgeons but
by medical men. Finally, it illustrates how, in stimulating such responses,
sick animals were able to shape medical knowledge and practice, and
how the zoo was run. Through exploring these themes in relation
3 Haughton (1864–1866).
4 Haughton (1864–1866, 1873), Adelman (2009).
5 de Courcy (2010).
2 DOCTORS IN THE ZOO: CONNECTING HUMAN AND ANIMAL HEALTH … 29
to the London and Dublin Zoological Gardens (established in 1828 and

1831, respectively), with occasional references to Bristol (1835) and
Manchester Belle Vue Zoological Gardens (1836), this chapter demon-
strates the mutual shaping of animals, medicine and zoological gardens in
the middle decades of the nineteenth century.
This period saw many zoological gardens established across Western
Europe. Symbolizing colonial possession and mastery over nature, they
were intended as bourgeois institutions, distinct from existing menag-
eries. Engaging in new modes of animal display, public education and
entertainment, zoos sought to advance knowledge of taxonomy, natu-
ral history, acclimatization, animal behaviour and comparative anatomy.
These aspects of their histories are well documented. However, the
health of their animal inhabitants is not.6
While animal historians have explored the lives and afterlives of cer-
tain zoo animals, and their contributions to human history, they have
paid little sustained attention to their health.7 This is surprising given
the extraordinarily high incidence of disease and death reported by
mid-nineteenth-century zoos. In London, for example, mortality rates
approached 33% per year,8 which suggests that ill health was fundamental
to the lived experiences of its animals. This chapter aims to shed light on
those experiences, and how animals were affected by human responses
to them. It thereby addresses issues neglected by zoo historians, who
are generally more concerned with the humans who founded, ran and
visited zoological gardens than the animals that lived within them. On
the occasions that these authors refer to animal health, they make ret-
rospective, negative assessments of human responses to it.9 This chap-
ter challenges such assessments by revealing the considerable attention
6 Green-Armytage (1964), Akerberg (2001), Keeling (2001), Baratay and Hardouin-
Fugier (2002), Burkhardt (2002), de Courcy (2009), Nyhart (2009) pp. 79–124, Ito
(2014).
7 Ritvo (1987) pp. 205–42, Rothfels (2002), Benbow (2004), Adelman (2009), Alberti
(2011), Flack (2013), Miller (2013), Flack (2014), Nance (2015).

8 Murie (1866).
9 For example: Akerberg (2001) pp. 186–94, Hancocks (2001) pp. 50–1, 73–6, Baratay
and Hardouin-Fugier (2002) pp. 131–9, Burt (2002), Cowie (2014) pp. 94–8. Many of
these authors rely on Chalmers Mitchell (1929), who as secretary of the Zoological Society
of London, 1903–1935, claimed to have taken the first real steps to improve zoo animal
health.
30 A. Woods
that diseased animals attracted during this period, and the many ways in
which medical men attempted to understand and promote their health.
The health of zoo animals is equally overlooked by medical historians.
As noted in Chapter 1, disciplinary traditions hold that medical history
is a field concerned primarily with human health, which considers ani-
mals only in their relations to humans. It positions sick animals within
the sub-field of veterinary history, and the study of animal life within the
history of biology. While these two fields have paid some attention to the
bodies and diseases of zoo animals,10 the absence of these animals from
medical history scholarship implies that their health had no bearing on
human lives. This chapter demonstrates to the contrary. It reveals doc-
tors’ efforts to advance zoo animal health for its own sake, how their
efforts intersected with veterinary practice and the study of compara-
tive anatomy, and how zoo animals contributed to knowledge of human
health. It thereby challenges historians’ very notions of medicine as a
human-focused endeavour.
The history recorded in this chapter derives from the traces that zoo
animals left on the medical historical record.11 These traces survive in
museum collections and catalogues, press reports, records of medi-
cal society meetings, medical journals and textbooks, medical biogra-
phies and the zoos’ institutional archives. They include the material
changes that disease inflicted on animal bodies, and their representa-
tion in images, verbal reports and statistics. They also encompass human
responses to those changes, which left imprints on human and animal
bodies and relationships, the zoos’ natural and built environments, and
on the careers of medical investigators. Through analysing these traces
and the circumstances of their production, the chapter sheds new light
on animals’ health histories, and on the historical co-constitution of ani-
mals, zoos and medicine.
The chapter is divided into halves. Each is structured around a dif-
ferent reason why zoo animal health attracted the attention of human
10 Veterinary accounts include: Jones (1976), Furman (1996). For the history of biol-
ogy, see Desmond (1985), Burkhardt (1999), Hochadel (2005), Hochadel (2011), Nyhart
(2009) pp. 110–7. Nyhart argues that a sense of moral obligation to animals provided an
important motivation for maintaining their health, but this was not evident in British zoos
at the time.
11 Benson (2011).
2 DOCTORS IN THE ZOO: CONNECTING HUMAN AND ANIMAL HEALTH … 31
doctors, and explores the interventions they made, and the implications
for animals and medicine. The first half revolves around the threat that
diseased animals posed to zoos as financially viable institutions devoted
to the scientific study of comparative anatomy. It recounts how medical
members of the zoological societies that ran the zoos attempted to pre-
vent, manage and learn about animal diseases through the use of three
modes of medicine that were typically applied to humans: public health,
bedside medicine and hospital medicine.12 Their use in the zoo awarded
diseased animals a quasi-human status, and refashioned them—in ways
shaped by the animals’ physical and behavioural characteristics—into vic-
tims of their environments, patients and pathological specimens, with
some unanticipated implications for human health.
The second half of the chapter explores the zoo’s appeal to medical
men who were not involved in its maintenance. This appeal lay in the
diversity of species, the presence of monkeys (whose zoological proximity
to humans was acknowledged long before Darwin) and the zoo’s status
as a total institution in which animal bodies, behaviours, lifestyles and
environments were centrally controlled by humans. Refashioning ani-
mals into points of comparison with humans, these doctors used them
to gather insights into human health, the general nature of disease, and
relationships between species.13 This agenda became known as ‘compara-
tive pathology’. Emerging at the nexus of medicine, veterinary medicine
and comparative anatomy, it was a quite different form of comparative
pathology to the experimental, laboratory-based comparative pathology
pursued by Pasteur, Koch and others, which dominates existing medical
historical literature.14 Like the health interventions documented in the
first half of the chapter, its analysis reveals that zoo animals exerted a far
greater influence on medical knowledge and practice than historians have
previously realized.
12 There is copious medical historical literature on these regimes. Key works include:
Foucault (1973), Jewson (1976), Hamlin (1998). For an overview, see Bynum (1994).
13 The concept of ‘total institution’ is usually attributed to Goffman, who described it
as ‘a place of residence and work where a large number of like-situated individuals, cut
off from the wider society for an appreciable period of time, together lead an enclosed,
formally administered round of life’ (Goffman 1968) p. 11. While this is a human-centred
definition, which Goffman applied to mental hospitals, it resonates with animal life in zoos.
14 Wilkinson (1992).
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26 „Proc. Zool. Soc.”, 1835, blz. 54. De zwartvleugel-pauw wordt door den heer Sclater als een bijzondere soort
beschouwd en heeft den naam van Pavo nigripennis ontvangen. ↑
27 Rudolphi, „Beiträge zur Anthropologie”, 1812, blz. 184. ↑
28 „Die Darwin’sche Theorie, und ihre Stellung zu Moral und Religion”, 1869, blz. 59. ↑
29 Dit wordt medegedeeld door den heer A. Leith Adams, in zijn „Field and Forest Rambles”, 1873, blz. 76, en stemt
overeen met zijn eigen ondervinding. ↑
30 Ten opzichte van de pauwen, zie Sir R. Heron, „Proc. Zoolog. Soc.”, 1835, blz. 54, en den weleerw. heer E. S.
Dixon, „Ornamental Poultry”, 1848, blz. 8. Omtrent den kalkoen, Audubon, ibid., blz. 4. Over den grooten auerhaan,
Lloyd, „Game Birds of Sweden”, blz. 23. ↑
31 De heer Hewitt, aangehaald in „Tegetmeier’s Poultry Book”, 1866, blz. 165. ↑
32 Aangehaald in Lloyd’s „Game Birds of Sweden”, blz. 345. ↑
33 Volgens Dr. Blasius („Ibis”, vol. II, 1860, blz. 297), zijn er 425 ontwijfelbare soorten van vogels die in Europa
broeien, behalve 60 vormen die veelvuldig als zelfstandige soorten worden beschouwd. Van deze laatste meent
Blasius, dat tien werkelijk twijfelachtig zijn, en dat de andere vijftig met hun naaste verwanten behooren te worden
vereenigd; maar dit bewijst, dat er een aanzienlijke mate van verscheidenheid bij sommige van onze Europeesche
vogels bestaat. Het is bij de natuuronderzoekers ook een nog niet uitgemaakt punt of verscheidene Noord-
Amerikaansche vogels moeten worden beschouwd als soortelijk onderscheiden van de overeenkomstige
Europeesche soorten. ↑
34 „Mammals and Birds of East Florida”, ook een „Ornithological Reconnaissance of Texas”, enz. Niettegenstaande
den invloed van het klimaat op de kleuren van vogels is het moeilijk de doffe of donkere tinten te verklaren van
bijna alle soorten die sommige landen bewonen, b.v. de Galapagoseilanden onder den evenaar, de uitgestrekte
gematigde vlakten van Patagonië, en, naar het schijnt, Egypte (zie den heer Hartshorne in „the American Naturalist”,
1873, blz. 747.) Deze landen zijn open en bieden den vogels weinig schuilplaatsen; maar het schijnt twijfelachtig of de
afwezigheid van levendig gekleurde soorten kan worden verklaard volgens het beginsel van bescherming, want op de
Pampa’s die ook open zijn, hoewel bedekt met groen gras, en waar de vogels evenzeer aan gevaar zijn blootgesteld,
zijn vele schitterende en opzichtig gekleurde soorten algemeen. Ik heb somtijds bespiegelingen gemaakt, of de
heerschende doffe tinten van het landschap in bovengenoemde landen geen invloed kunnen hebben gehad op het
waarnemingsvermogen voor kleuren van de vogels die ze bewonen. ↑
35 „Ontstaan der Soorten”, 3de Ned. uitgaaf, blz. 129. Ik had altijd begrepen, dat zeldzame en sterke afwijkingen in
maaksel, die den naam van monstruositeiten verdienden, slechts zelden door natuurlijke teeltkeus konden bewaard
blijven, en dat het bewaard blijven zelfs van in hooge mate voordeelige afwijkingen tot op zekere hoogte van het
toeval af zou hangen. Ik had ook de belangrijkheid van individueele verschillen volkomen naar waarde geschat, en dit
bracht mij er toe om zoo sterk te drukken op de belangrijkheid van dien onbewusten vorm van teeltkeus door den
mensch, die het gevolg is van het bewaren van de hoogst geschatte individu’s van elk ras, zonder eenig oogmerk van
zijn kant om de kenmerken van het ras te wijzigen. Doch voordat ik een uitnemend artikel in de „North British Review”
[121](Maart 1867, blz. 589 v.v.), die mij van meer dienst is geweest dan eenig ander Review, had gelezen, zag ik niet
in, hoe groot de kansen waren tegen het bewaard blijven van afwijkingen, hetzij kleine of sterk uitgedrukte, die alleen
bij enkele individu’s voorkwamen. ↑
36 „Introduct. to the Trochilidae”, blz. 102. ↑
37 Gould, „Handbook of Birds of Australia”, vol. II, blz. 32 en 68. ↑
38 Audubon, „Ornitholog. Biography”, 1838, vol. IV, blz. 389. ↑
39 Jerdon, „Birds of India”, vol. I, blz. 108, en de heer Blyth in „Land and Water”, 1868, blz. 381. ↑
40 Graba, „Tagebuch, Reise nach Farö”, 1830, blz. 41–44. Macgillivray, „Hist. British Birds”, vol. III, blz. 745. „Ibis”, vol.
V, 1863, blz. 469. ↑
41 Graba, ibid., blz. 54. Macgillivray, ibid., vol. V, blz. 327. ↑
42 „Het Varieeren der Huisdieren en Cultuurplanten”, Ned. vert., deel II, blz. 55. ↑
43 Zie omtrent deze punten ook „Het Varieeren der Huisdieren en Cultuurplanten”, Ned. vert., deel I, blz. 295; deel II,
blz. 54. ↑
44 Zie b.v. over de regenboogvliezen (irides) van een Podica en Gallicrex in „Ibis”, vol. II, 1860, blz. 206; en vol. V,
1863, blz. 426. ↑
45 Zie ook Jerdon, „Birds of India”, vol. I, blz. 243–245. ↑
46 „Zoology of the Voyage of H. M. S. Beagle”, 1841, blz. 6. ↑
47 Bechstein, „Naturgeschichte Deutschlands”, Bd. IV, 1795, blz. 31, over een onder-verscheidenheid (sub-variëteit)
van de Monck-duif. ↑
48 Deze houtsnede is gegraveerd naar een fraaie teekening welke de heer Trimen de bijzondere vriendelijkheid had
voor mij te vervaardigen; zie ook [128]zijn beschrijving van de wonderlijke grootte der afwijking in de kleur en den
vorm van de vleugels van deze kapel, in zijn „Rhopalocera Africae Australis”, blz. 186. Zie ook een belangwekkende
verhandeling van den weleerw. heer H. H. Higgens over den oorsprong der oogvlekken bij de Schubvleugeligen
(Lepidoptera) in het „Quarterly Journal of Science”, Juli, 1860, blz. 325. ↑
49 Jerdon, „Birds of India”, vol. III, blz. 517. ↑
50 „Varieeren der Huisdieren en Cultuurplanten”, Ned. vert., deel I, blz. 294. ↑
51 Als de Argus-fazant zijn vleugelslagpennen als een grooten waaier tentoonspreidt, staan die welke het dichtst bij
het lichaam staan, meer rechtop dan de buitenste, zoodat de schaduwing van de bal-en-holte oogvlekken een
weinig verschillend behoorde te zijn op de verschillende vederen om haar vol effect met betrekking tot den inval van
het licht voort te brengen. De [138]heer T. W. Wood die het ervaren oog van een kunstenaar bezit, verzekert („Field”
Courant, 28 Mei 1870, blz. 457), dat dit het geval is; maar na zorgvuldig twee opgezette voorwerpen te hebben
onderzocht (van een waarvan ik de bedoelde vederen tot meer nauwkeurige vergelijking aan den heer Gould heb
gegeven), kan ik niet bemerken, dat dit toppunt van volmaaktheid in de schaduwing is bereikt, en evenmin hebben
anderen aan wie ik deze vederen heb getoond, dit feit kunnen opmerken. ↑
52 „The Field”, 28 Mei, 1870. ↑
53 „Popular Lectures on Scientific Subjects”, Eng. vert., 1873, blz. 219, 227, 269, 390. ↑
54 „The Reign of Law”, 1867, blz. 347. ↑
55 „The Naturalist in Nicaragua”, 1874, blz. 112. ↑
56 „Introduction to the Trochilidae”, 1861, blz. 110. ↑
57 „Zur Zeichnung der Vogelfedern”, Zeitschr. für wiss. Zoologie, Bd. XLIV. ↑
58 Met de patrijzen verwante vogels uit Zuid- en West-Azië en Afrika. ↑
59 Ceriornis Satyra, een tot de zoogenaamde hoornfazanten behoorende vogel uit de Himalaya en Zuid-China. ↑
[Inhoud]
VIJFTIENDE HOOFDSTUK.
SECUNDAIRE SEKSUEELE KENMERKEN BIJ DE
VOGELS.—VERVOLG.
Onderzoek, waarom bij sommige soorten alleen de mannetjes, en bij

andere beide seksen schitterende kleuren vertoonen.—Over tot ééne
sekse beperkte erfelijkheid: toepassing daarvan op verschillende deelen
en op een schitterend gekleurd vederkleed.—Betrekking tusschen
nestbouw en kleur.—Verlies van het bruiloftskleed gedurende den winter.
Wij moeten in dit hoofdstuk onderzoeken, waarom bij vele soorten

van vogels het wijfje niet de zelfde versierselen heeft ontvangen als
het mannetje; en waarom bij vele andere beide seksen de zelfde, of
bijna de zelfde kleuren vertoonen? In het volgende hoofdstuk zullen
wij onderzoeken, waarom in enkele zeldzame gevallen het wijfje
schitterender is dan het mannetje.
In mijn „Ontstaan der Soorten” 1 heb ik kortelijk het denkbeeld

geopperd, dat de lange staart van den pauw lastig en de in ’t oog
vallende zwarte kleur van den auerhaan gevaarlijk zouden zijn voor
het wijfje gedurende den broeitijd, en dat derhalve de overplanting
van deze kenmerken van het mannetje op zijn vrouwelijke
nakomelingschap door de natuurlijke teeltkeus was verhinderd. Het
is nog mijn meening, dat dit in eenige weinige gevallen kan zijn
gebeurd: doch na rijp nadenken over al de feiten die ik in staat ben
geweest te verzamelen, ben ik nu geneigd te gelooven, dat, wanneer
de seksen verschillen, de opeenvolgende afwijkingen over het
algemeen van den beginne af in haar overplanting beperkt zijn
geweest tot de zelfde sekse bij welke zij het eerst verschenen. Sinds
mijn opmerkingen in het licht verschenen, is het onderwerp van
seksueele kleuring in eenige hoogst belangwekkende
verhandelingen besproken door den heer Wallace 2, die gelooft, dat
in [150]bijna alle gevallen de opeenvolgende afwijkingen een neiging
bezaten om gelijkelijk op beide seksen te worden overgeplant; maar
dat het wijfje door de natuurlijke teeltkeus werd verhinderd om de
opzichtige kleuren van het mannetje te verkrijgen, ten gevolge van
het gevaar dat zij daardoor gedurende den broeitijd zou hebben
geloopen.
Deze zienswijze maakt een vervelende redekaveling over een

moeilijk punt noodzakelijk, namelijk of de overplanting van een
kenmerk dat eerst door beide seksen wordt overgeërfd, later in zijn
overplanting door middel der teeltkeus alleen tot ééne sekse kan
worden beperkt. Wij moeten bedenken, dat, gelijk in het inleidende
hoofdstuk over seksueele teeltkeus is aangetoond, kenmerken die in
hun ontwikkeling tot ééne sekse zijn beperkt, altijd bij de andere in
latenten toestand bestaan. Een denkbeeldig voorbeeld zal ons het
best helpen om de moeilijkheid van het geval te zien; laten wij eens
veronderstellen, dat een duivenfokker een duivenras wenscht te
vormen, bij hetwelk alleen de mannetjes bleek blauw zouden zijn
gekleurd, terwijl de wijfjes haar vroegere leiachtige kleur behielden.
Daar bij duiven kenmerken van alle soorten gewoonlijk gelijkelijk op
beide seksen worden overgeplant, zou de fokker moeten beproeven
dezen laatsten vorm van erfelijkheid in tot ééne sekse beperkte
overplanting te veranderen. Al wat hij kon doen, zou zijn om
voortdurend elke mannelijke duif die in de minste mate van een
bleeker blauwe kleur was, voor de voortteling uit te kiezen; en het
natuurlijk gevolg van deze handelwijze, als zij gedurende langen tijd
onophoudelijk werd voortgezet, en indien de bleeke afwijkingen sterk
werden overgeërfd of dikwijls terugkwamen, zou zijn om het geheele
geslacht van een lichter blauw te maken. Onze fokker zou echter
genoodzaakt zijn om in elke opeenvolgende generatie zijn
bleekblauwe mannetjes met leikleurige wijfjes te doen paren; want
hij wenscht, dat de laatste deze kleur behouden. De uitslag zou over
het algemeen zijn, hetzij de voortbrenging van een partij gevlekte
bastaarden, of wel nog waarschijnlijker het spoedig en volkomen
verloren gaan van de bleek-blauwe kleur; want de oorspronkelijke
leikleurige tint zou met overwegende kracht worden overgeplant.
Onderstellende echter, dat in elke opeenvolgende generatie enkele
bleek-blauwe mannetjes en leikleurige wijfjes werden voortgebracht,
en dat deze altijd met elkander werden gepaard, dan zouden de
leikleurige wijfjes, als ik de uitdrukking mag gebruiken, veel blauw
bloed in haar aderen hebben; want haar vaders, grootvaders enz.
zouden blauwe vogels zijn geweest. Onder deze omstandigheden
[151]zou het te begrijpen zijn (hoewel ik geen stellige feiten ken, die
het waarschijnlijk maken), zoo de leikleurige wijfjes een zoo groote
latente neiging tot bleek-blauwheid verkregen, dat zij die kleur bij
haar mannelijke nakomelingschap niet vernietigden, terwijl haar
vrouwelijke nakomelingschap de leikleurige tint bleef behouden.
Indien dit zoo was, zou het begeerde einddoel om een ras te maken,
bij hetwelk de beide seksen standvastig in kleur verschillen, kunnen
worden bereikt.
De uiterste belangrijkheid of liever noodzakelijkheid, dat in het

bovengenoemde geval het gewenschte kenmerk, namelijk bleek-
blauwheid, bij het wijfje, hoewel in latenten staat, bestond, zal het
best door het volgende voorbeeld worden gewaardeerd: het
mannetje van den Soemmerring’s fazant heeft een staart van meer
dan 90 centimeter lengte, terwijl die van het wijfje slechts ruim 20
centimeter lang is; de staart van het mannetje van den gewonen
fazant is omstreeks 50 centimeter lang, en die van zijn wijfje
omstreeks 30 centimeter. Indien nu het wijfje van den
Soemmerring’s fazant met haar korten staart met het mannetje van
den gewonen fazant werd gekruist, kan er geen twijfel zijn, of het
mannelijke bastaardkroost zou een veel langeren staart hebben dan
die van het onvermengde kroost van den gewonen fazant. Indien
daarentegen het wijfje van den gewonen fazant, met haar staart
welke bijna tweemaal zoo lang (1) als die van het wijfje van den
Soemmerring’s fazant is, met het mannetje van dezen laatsten werd
gekruist, zou het mannelijke bastaardkroost een veel korter staart
hebben dan die van het onvermengde kroost van den
Soemmerring’s fazant. 3
Om zijn nieuw ras met mannetjes van een beslist bleek-grauwe kleur
en onveranderde wijfjes te maken, zou onze fokker gedurende vele
generatiën voort moeten gaan met de mannetjes voor de voortteling
uit te kiezen; en elke graad van bleekheid zou bij de mannetjes
moeten worden gefixeerd en bij de wijfjes latent gemaakt. Dit zou
een uiterst moeilijke taak zijn en is nimmer beproefd, maar zou
mogelijk slagen. De voornaamste hinderpaal zou het spoedige en
volkomen verlies van de bleek-blauwe kleur zijn, wegens de
noodzakelijkheid van herhaalde [152]kruisingen met het leikleurige
wijfje, daar dit laatste in het eerst volstrekt geen latente neiging bezit,
om bleekblauw kroost voort te brengen.
Indien daarentegen een of twee mannetjes in hoe geringe mate ook

in bleekheid afweken, en de afwijkingen van den beginne af in haar
overplanting tot de mannelijke sekse beperkt bleven, zou de taak om
een nieuw ras van de begeerde soort te maken, gemakkelijk zijn;
want men zou eenvoudig de mannetjes voor de voortteling hebben
uit te kiezen en hen met gewone wijfjes te paren. Een soortgelijk
geval heeft werkelijk plaats gehad; want er zijn in België 4
duivenrassen bij welke alleen de mannetjes met zwarte strepen zijn
geteekend. Zoo heeft ook de heer Tegetmeier voor eenige jaren
aangetoond 5, dat Engelsche Pagadet-duiven („dragons”) niet zelden
zilverkleurige jongen voortbrengen, dat gewoonlijk wijfjes zijn, en hij
heeft zelf tien zulke wijfjes uitgebroeid. Daarentegen is het hoogst
zeldzaam, dat er een zilverkleurig mannetje wordt voortgebracht,
zoodat niets gemakkelijker is dan een ras van Pagadet-duiven
(„dragons”) te vormen, waarbij de mannetjes blauw en de wijfjes
zilverkleurig zijn. Deze neiging is inderdaad zoo sterk, dat, toen de
heer Tegetmeier ten laatste een zilverkleurig mannetje verkreeg en
dat met een zilverkleurig wijfje paarde, hij in zijn verwachting om een
ras te verkrijgen, waarin beide seksen die kleur bezaten, werd
teleurgesteld; want het jonge mannetje keerde terug tot de blauwe
kleur van zijn grootvader, en alleen het jonge wijfje was zilverkleurig.
Ongetwijfeld zou, als men geduld gebruikte, deze neiging tot
atavisme bij de mannetjes, gesproten uit de vereeniging van een
toevallig zilverkleurigen doffer met een zilverkleurige duif, kunnen
worden vernietigd, en dan zouden beide seksen gelijk zijn gekleurd;
en juist deze handelwijze is door den heer Esquilant met goed
gevolg aangewend in het geval van zilverkleurige meeuwtjes
(„turbits”). Bij hoenders komen afwijkingen in kleur, die in haar
overplanting tot de mannelijke sekse beperkt zijn, veelvuldig voor.
Zelfs als deze vorm van erfelijkheid de overhand behield, zou het
wel kunnen gebeuren, dat sommige van de opeenvolgende stappen
in het proces van afwijking op het wijfje werden overgebracht, dat er
dan toe zou komen om eenigermate op het mannetje te gelijken,
zooals bij sommige hoenderrassen het geval is. Of ook het grootste
aantal, maar niet alle opeenvolgende stappen zouden op beide
seksen kunnen worden overgebracht, en het [153]wijfje zou dan zeer
veel op het mannetje gelijken. Het kan nauwelijks worden betwijfeld,
dat dit de oorzaak is, waarom het mannetje van de Kropduif een iets
grooteren kop en dat van de Postduif iets grootere vleeschlappen
hebben dan hun respectieve wijfjes; want de fokkers hebben niet
van de eene sekse met meer zorg individu’s voor de voortteling
uitgezocht dan van de andere, en hebben den wensch niet
gekoesterd, dat het mannetje in hooger mate met deze kenmerken
zou prijken dan het wijfje, en toch is dit bij beide rassen het geval.
De zelfde handelwijze zou moeten worden gevolgd, en men zou de

zelfde moeilijkheid ontmoeten, wanneer men een ras wenschte te
vormen, waarvan alleen de wijfjes van de eene of andere nieuwe
kleur waren. Onze fokker zou eindelijk een ras kunnen wenschen te
vormen, waarbij de twee seksen van elkander en beide van de
stamsoort verschilden. Hier zou de moeilijkheid uiterst groot zijn,
tenzij de opeenvolgende afwijkingen van het begin af van beide
seksen tot eene sekse waren beperkt, en dan zou er geen
moeilijkheid bestaan. Wij zien dat bij de Hoenders; zoo verschillen
de beide seksen van de gepenseelde Hamburger hoenders zeer van
elkander, zoowel als van de beide seksen van den oorspronkelijken
Gallus bankiva; en beide worden nu bestendig op haar standaard
van uitnemendheid gehouden door voortgezette teeltkeus, hetgeen
onmogelijk zou zijn, wanneer niet de onderscheidene kenmerken
van beide in hun overplanting waren beperkt. De Spaansche
hoenders bieden een merkwaardig geval aan; de haan bezit een
verbazend grooten kam, maar sommige van de opeenvolgende
afwijkingen, door de opeenhooping waarvan hij dien heeft verkregen,
schijnen op de hen te zijn overgebracht; want zij heeft een kam die
vele malen grooter is dan die van de hennen van de stamsoort. De
kam van de hen verschilt echter in één opzicht van dien van den
haan, want hij is geneigd om over te hangen, en in den laatsten tijd
heeft de mode beslist, dat dit altijd het geval behoorde te zijn, en dit
bevel is spoedig met goeden uitslag gevolgd. Nu moet het
overhangen van den kam wel in zijn overplanting seksueel beperkt
zijn; want anders zou het verhinderen, dat de kam van den haan
volkomen rechtstandig bleef hetgeen in het oog van elken fokker
afschuwelijk zou zijn. Daarentegen moet ook de rechtstandigheid
van den kam van den haan eveneens een seksueel beperkt
kenmerk zijn; want anders zou het het overhangen van den kam van
de hen tegengaan.
Uit de voorgaande voorbeelden zien wij, dat het, zelfs als men een
[154]bijna onbegrensden tijd tot zijn beschikking had, een uiterst
moeilijk en ingewikkeld, hoewel wellicht niet onmogelijk proces zou
zijn om door teeltkeus den eenen vorm van erfelijkheid in den
anderen te veranderen. Daarom ben ik, zonder stellige bewijzen in
elk afzonderlijk geval, ongeneigd om aan te nemen, dat dit bij
natuurlijke soorten dikwijls is geschied. Daarentegen zou er, door
middel van opeenvolgende afwijkingen die van den beginne af in
haar overplanting seksueel waren beperkt, geen de minste
moeilijkheid in zijn gelegen om een mannelijken vogel in kleur of in
eenig ander kenmerk zeer verschillend van het wijfje te maken,
terwijl dit laatste onveranderd bleef, of slechts weinig veranderd, of
bijzonder ter wille van de bescherming werd gewijzigd.
Daar levendige kleuren aan de mannetjes van dienst zijn bij hun
mededinging met hun medeminnaars, zullen dergelijke kleuren voor
de voortteling worden uitgezocht, hetzij zij al dan niet uitsluitend op
de zelfde sekse worden overgeplant. Men zou bijgevolg mogen
verwachten, dat de wijfjes dikwijls in meerdere of mindere mate in de
levendige kleuren van de mannetjes zouden deelen; en dit is
werkelijk bij een menigte soorten het geval. Indien al de
opeenvolgende afwijkingen gelijkelijk op beide seksen werden
overgeplant, zouden de wijfjes niet van de mannetjes zijn te
onderscheiden; en dit is eveneens bij vele vogels het geval. Indien
echter doffe kleuren van hoog belang waren voor de veiligheid van
het wijfje gedurende den broeitijd, gelijk bij vele op den grond
nestelende vogels, zouden de wijfjes die door levendige kleuren
afweken of door overerving van de mannetjes eenige merkbare
toeneming van de levendigheid hunner kleuren verkregen, vroeger
of later te gronde gaan. De neiging in de mannetjes om gedurende
een onbeperkten tijd voort te gaan met op hun vrouwelijke
nakomelingen hun eigen levendigheid van kleur over te planten, zou
moeten worden geëlimineerd door een verandering in den vorm van
erfelijkheid; en dit zou, gelijk door ons voorafgaand voorbeeld wordt
aangetoond, uiterst moeilijk zijn. Het meer waarschijnlijk gevolg van
de lang voortgezette vernieling van de meer levendig gekleurde
wijfjes, veronderstellende dat de gelijke vorm van overplanting de
overhand behield, zou zijn de vermindering of vernietiging van de
levendige kleuren van de mannetjes, ten gevolge van hun
voortdurende kruising met de doffer gekleurde wijfjes. Het zou
vervelend zijn alle andere mogelijke gevolgen ten einde toe na te
gaan; ik mag echter den lezer herinneren dat, gelijk in het achtste
hoofdstuk is aangetoond, indien [155]zich bij de wijfjes seksueel
beperkte afwijkingen in levendigheid van kleur voordeden, deze,
zelfs al waren zij in het minst niet nadeelig voor hen en al werden zij
bij gevolg niet geëlimineerd, toch niet zouden worden begunstigd of
voor de voortteling uitgekozen; want het mannetje neemt gewoonlijk
elk wijfje aan en kiest de meer aantrekkelijke individu’s niet voor de
voortteling uit; bij gevolg zouden deze afwijkingen er aan zijn
blootgesteld verloren te gaan, en weinig invloed hebben op de
kenmerken van het ras; en dit zal helpen om te verklaren, waarom
de wijfjes gewoonlijk minder levendig zijn gekleurd dan de
mannetjes.
In het juist aangehaalde hoofdstuk werden voorbeelden gegeven,

die, zooveel men maar wilde, zouden kunnen worden vermeerderd,
van afwijkingen die zich op verschillende leeftijden voordeden en op
die zelfde leeftijden werden overgeërfd. Er werd ook aangetoond,
dat afwijkingen die zich laat in het leven voordoen, gewoonlijk
worden overgeplant op de zelfde sekse bij welke zij het eerst
verschenen, terwijl afwijkingen die zich vroeg in het leven voordoen,
geneigd zijn om op beide seksen te worden overgeplant, hoewel alle
gevallen van seksueel beperkte overplanting niet op die wijze
kunnen worden verklaard. Verder werd aangetoond, dat, wanneer
een mannelijke vogel afweek door levendiger te worden gekleurd,
terwijl hij jong was, dergelijke afwijkingen van geen dienst zouden
zijn, voordat de leeftijd was gekomen, waarop hij zich voortplantte,
en er wedstrijd was tusschen mannetjes die elkanders
medeminnaars waren. In het geval van vogels die op den grond
leven en gewoonlijk de bescherming van doffe kleuren noodig
hebben, zouden echter levendige kleuren veel gevaarlijker voor de
jonge en nog geen ondervinding hebbende dan voor de volwassen
mannetjes zijn. Bij gevolg zouden de mannetjes die door
levendigheid van kleur afweken, terwijl zij jong waren, aan veel
vernieling lijden en door natuurlijke teeltkeus worden geëlimineerd;
daarentegen zouden de mannetjes die op die wijze afweken, als zij
omtrent volwassen waren, niettegenstaande zij aan een weinig meer
gevaar waren blootgesteld, kunnen blijven leven, en daar zij door de
seksueele teeltkeus waren begunstigd, hun soort voortplanten. Het
vernietigd worden van de levendig gekleurde jonge mannetjes en het
voorspoedig zijn der volwassenen in hun vrijage kan, volgens het
beginsel, dat er een betrekking bestaat tusschen het levenstijdperk
waarin de afwijking plaats heeft, en den vorm van overplanting,
verklaren, dat van vele vogels [156]alleen de mannetjes schitterende
kleuren hebben verkregen en die alleen op hun mannelijke
nakomelingschap hebben overgeplant. Ik wensch echter in geenen
deele vol te houden, dat de invloed van den leeftijd op den vorm van
overplanting indirect de eenige oorzaak is van het groote verschil in
de pracht van het gevederte tusschen de seksen van vogels.
Daar het bij alle vogels bij welke de seksen in kleur verschillen, een
belangwekkende vraag is, of alleen de mannetjes door seksueele
teeltkeus zijn gewijzigd, en de wijfjes, zoover de werking van dit
beginsel aangaat, onveranderd of bijna onveranderd zijn gelaten;
dan wel of de wijfjes bijzonder zijn gewijzigd door natuurlijke
teeltkeus ter wille van de bescherming, zal ik dit vraagstuk
uitvoeriger bespreken, uitvoeriger zelfs dan zijn innerlijke
belangrijkheid verdient; want onderscheidene merkwaardige
daarmede zijdelings in verband staande punten kunnen dan tevens
gepast worden beschouwd.
Voor wij een aanvang maken met het onderwerp van de kleur, meer
bijzonder in verband met de besluiten van den heer Wallace, kan het
wellicht nuttig zijn uit een gelijksoortig oogpunt eenige andere
verschillen tusschen de seksen te beschouwen. Vroeger bestond er
een ras van hoenders in Duitschland 6, bij hetwelk de hennen sporen
bezaten; zij waren goede eierlegsters; maar zij brachten haar nesten
met haar sporen zoozeer in de war, dat men ze haar eigen eieren
niet kon laten uitbroeien. Van daar was er een tijd, dat het mij
waarschijnlijk toescheen, dat bij de wijfjes van de wilde
Hoenderachtige Vogels (Gallinaceae) de ontwikkeling van sporen
door de natuurlijke teeltkeus was verhinderd, wegens de daardoor
aan de nesten toegebrachte schade. Dit scheen des te
waarschijnlijker, omdat de vleugelsporen die het nest geen schade
konden berokkenen, dikwijls bij het wijfje even goed waren
ontwikkeld als bij het mannetje, hoewel zij in niet weinig gevallen bij
het mannetje iets grooter zijn. Als het mannetje sporen aan de
pooten bezit, vertoont het wijfje bijna altijd rudimenten daarvan,—het
rudiment bestaat somtijds eenvoudig uit een schub, zooals bij de
soorten van het geslacht Gallus. Men zou daarom kunnen beweren,
dat de wijfjes oorspronkelijk goed ontwikkelde sporen hadden
bezeten, maar dat zij deze later hadden verloren, hetzij door onbruik
of door natuurlijke teeltkeus. Indien deze beschouwingswijze echter
werd aangenomen, zou zij moeten worden [157]uitgebreid tot tallooze
andere gevallen; en zij sluit in zich, dat de vrouwelijke voorouders
van de bestaande spoordragende soorten eens waren overladen
met een nadeelig aanhangsel.
Bij eenige weinige geslachten en soorten, zooals bij Galloperdix,

Acomus, en den Javaanschen pauw (Pavo muticus), bezitten zoowel
de wijfjes als de mannetjes goed ontwikkelde sporen. Moeten wij uit
dit feit afleiden, dat zij een soort van nest bouwen, niet vatbaar om
door hun sporen te worden beschadigd, en verschillend van dat van
hun naaste verwanten, zoodat hier geen noodzakelijkheid bestond
om de sporen te doen verdwijnen? Of moeten wij veronderstellen,
dat deze wijfjes bijzonder behoefte aan sporen hebben voor haar
verdediging? Het is een meer waarschijnlijk besluit, dat zoowel de
aanwezigheid als de afwezigheid van sporen bij de wijfjes een
gevolg zijn van het de overhand behouden van verschillende wetten
van overerving, onafhankelijk van de natuurlijke teeltkeus. Omtrent
de vele wijfjes bij welke zich sporen in rudimentairen toestand
vertoonen, mogen wij besluiten, dat eenige weinige van de
opeenvolgende afwijkingen door welke zij zich bij de mannetjes
ontwikkelden, zich vroeg in het leven vertoonden, en ten gevolge
daarvan op de wijfjes werden overgeplant. In de andere en veel
zeldzamer gevallen in welke de wijfjes volkomen ontwikkelde sporen
bezitten, mogen wij besluiten, dat al de opeenvolgende afwijkingen
op haar werden overgebracht, en dat zij trapsgewijze de overgeërfde
gewoonte verkregen om haar nesten niet in de war te brengen.
De stemorganen en de op verschillende wijzen om geluid voort te

brengen gewijzigde vederen, zoowel als de eigenaardige instinkten
om ze te gebruiken, verschillen dikwijls bij de twee seksen, maar zijn
somtijds bij beide de zelfde. Kan men dergelijke verschijnselen
verklaren, doordat de mannetjes deze organen en instinkten hebben
verkregen, terwijl de wijfjes zijn verhinderd om ze over te erven, ten
gevolge van het gevaar waaraan zij blootgesteld zouden zijn
geweest door de aandacht van roofvogels of roofdieren tot zich te
trekken? Dit schijnt mij niet waarschijnlijk, wanneer wij denken aan
de menigte vogels die gedurende de lente straffeloos het land met
hun stem opvroolijken. 7 Het is een veiliger besluit, dat, daar vocale
en instrumentale organen [158]alleen van dienst zijn aan de
mannetjes gedurende hun vrijage, deze organen alleen bij deze
sekse door seksueele teeltkeus en voortdurend gebruik tot
ontwikkeling kwamen,—terwijl de opeenvolgende afwijkingen en de
gevolgen van het gebruik van den beginne af in hun overplanting in
meerdere of mindere mate alleen tot de mannelijke sekse beperkt
bleven.
Vele soortgelijke gevallen zouden kunnen worden aangevoerd, bijv.
de vederen op den kop, die over het algemeen bij het mannetje
langer zijn dan bij het wijfje, somtijds bij beide seksen even lang zijn,
en nu en dan bij het wijfje ontbreken,—terwijl deze verschillende
gevallen dikwijls in ééne en de zelfde groep vogels worden
aangetroffen. Het zou moeilijk zijn een verschil van deze soort
tusschen de seksen te verklaren volgens het beginsel, dat het wijfje
was bevoordeeld door het bezit van een weinig korter kuif dan het
mannetje en het ten gevolge daarvan kleiner worden of volkomen
verdwijnen van die kuif door natuurlijke teeltkeus. Ik zal echter een
gunstiger geval nemen, namelijk de lengte van den staart. De lange
staart van den pauw zou niet slechts lastig, maar zelfs gevaarlijk zijn
geweest voor de pauwin gedurende den broeitijd en terwijl zij haar
jongen vergezelt. Daarom is het a priori in het minst niet
onwaarschijnlijk, dat de ontwikkeling van haar staart door natuurlijke
teeltkeus is belet. De wijfjes van onderscheidene fazanten die in
haar open nesten blijkbaar aan evenveel gevaar zijn blootgesteld
geweest als de pauwin, hebben echter staarten van aanmerkelijke
lengte. De wijfjes van den Liervogel (Menura superba) hebben
evengoed lange staarten als de mannetjes, en zij bouwen
koepelvormige nesten, hetgeen bij zulk een grooten vogel een
groote afwijking is. De natuuronderzoekers zijn er verwonderd over
geweest, hoe het wijfje van den Liervogel gedurende het broeien
met haar staart kon klaar komen; men weet nu echter 8, dat zij „eerst
haar kop in het nest steekt, en dan ronddraait, haar staart somtijds
over den rug, maar meer veelvuldig langs haar zijde omgebogen
houdende. De staart wordt daardoor na eenigen tijd geheel scheef,
en is een vrij bruikbare aanwijzing van de lengte van tijd, gedurende
welken de vogel op haar eieren heeft gezeten.” Bij beide seksen van
een Australischen IJsvogel (Thanysiptera sylvia) zijn de middelste
staartvederen zeer lang; en daar het wijfje haar nest in een gat
maakt, worden deze [159]vederen, gelijk de heer R. B. Sharpe mij
meldt gedurende den nestbouw zeer verfrommeld.
In deze beide gevallen moet de groote lengte der staartvederen
eenigermate lastig voor het wijfje zijn; en daar bij beide soorten de
staartvederen van het wijfje iets korter zijn dan die van het mannetje,
zou men kunnen beweren, dat hun volkomen ontwikkeling door de
natuurlijke teeltkeus was belet. Te oordeelen naar deze gevallen,
zou de pauwin, wanneer de ontwikkeling van haar staart alleen was
verhinderd, toen hij lastig of gevaarlijk lang werd, een veel langeren
staart hebben verkregen dan zij werkelijk bezit; want haar staart is
op verre na zoo lang niet, in verhouding tot de grootte van haar
lichaam, als die van vele vrouwelijke fazanten, en ook niet langer
dan die van de kalkoensche hen. Men moet ook steeds bedenken,
dat, zoodra als in overeenstemming met deze beschouwingswijze de
staart van de pauwin gevaarlijk lang werd en haar ontwikkeling bij
gevolg werd verhinderd, dit voortdurend zou hebben teruggewerkt
op haar mannelijke nakomelingschap, en dus den pauw zou hebben
belet om zijn tegenwoordigen prachtigen staart te verkrijgen. Wij
mogen daarom de gevolgtrekking maken, dat de lengte van den
staart bij den pauw en zijn kortheid bij de pauwin zijn veroorzaakt,
doordat de vereischte afwijkingen van het mannetje van den beginne
af alleen op de mannelijke nakomelingen zijn overgeplant.
Wij worden tot omtrent het zelfde besluit gebracht ten opzichte van
de lengte van den staart bij de onderscheidene soorten van
fazanten. Bij den geoorden fazant (Crossoptilon auritum) is de staart
bij beide seksen even lang, namelijk 40 of 42,5 centimeter; bij den
gewonen fazant is hij bij het mannetje omtrent 50 centimeter, en bij
het wijfje 30 centimeter lang; bij Soemmerring’s fazant 92,5
centimeter bij het mannetje en slechts 20 bij het wijfje; en bij Reeve’s
fazant eindelijk is hij werkelijk bij het mannetje soms 180 centimeter
en bij het wijfje 40 centimeter lang. Bij de verschillende soorten
verschilt dus de staart van het wijfje veel in lengte, en wel niet in
verhouding van de lengte van den staart bij de respectieve
mannetjes der zelfde soorten; en dit kan, naar het mij toeschijnt, met
veel meer waarschijnlijkheid worden verklaard door de wetten der
erfelijkheid,—dat is doordat de opeenvolgende afwijkingen van den
beginne af in haar overplanting meer of minder volkomen beperkt
zijn gebleven tot de mannelijke sekse,—dan door de werking der
natuurlijke teeltkeus, die het gevolg zou zijn [160]geweest van het
nadeel dat de lengte van den staart in meerdere of mindere mate
aan de wijfjes der verschillende soorten berokkende.
Wij kunnen nu overgaan tot de beschouwing van de bewijsgronden

van den heer Wallace ten opzichte van de seksueele kleuring van
vogels. Hij gelooft, dat de levendige kleuren, oorspronkelijk door de
mannetjes door seksueele teeltkeus verkregen, in alle of bijna alle
gevallen op de wijfjes zouden zijn overgebracht, wanneer de
overplanting niet door de natuurlijke teeltkeus ware verhinderd. Ik
herinner hier den lezer, dat onderscheidene feiten, op deze meening
betrekking hebbende, bij de behandeling der Reptielen, Amphibiën,
Visschen en Schubvleugelige Insekten zijn medegedeeld. De heer
Wallace steunt die meening hoofdzakelijk, maar niet uitsluitend,
gelijk wij in het volgende hoofdstuk zullen zien, op de volgende
mededeeling 9, dat, wanneer beide seksen op sterk opzichtige wijze
zijn gekleurd, het nest van zoodanigen aard is, dat het den op de
eieren zittenden vogel verbergt; maar dat, wanneer er een sterk
uitgedrukt verschil van kleur tusschen de seksen bestaat, zoodat het
mannetje levendig en het wijfje dof is gekleurd, het nest open is en
den op de eieren zittenden vogel aan het gezicht blootstelt. Deze
overeenstemming, zoover zij gaat, steunt ongetwijfeld het geloof, dat
de wijfjes die op open nesten zitten, bijzonder zijn gewijzigd ter wille
van de bescherming. De heer Wallace geeft toe, dat er, gelijk kon
worden verwacht, eenige uitzonderingen op zijn beide regels
bestaan; het is echter de vraag of deze uitzonderingen niet zoo talrijk
zijn, dat zij die ernstig verzwakken.
Er is in de eerste plaats veel waars in de opmerking van den Hertog
van Argyll 10, dat een groot koepelvormig nest gemakkelijker in het
oog valt aan den vijand, vooral aan alle op boomen verblijf houdende
roofdieren, dan een kleiner open nest. Wij moeten ook niet vergeten,
dat bij vele vogels die open nesten bouwen, de mannetjes op de
eieren zitten en in het voeden der jongen behulpzaam zijn even
goed als de wijfjes; dit is bij voorbeeld het geval bij den zomer-
roodvogel of vuurtanagra (Pyranga aestiva) 11, een der prachtigste
vogels van de Vereenigde Staten, waarvan het mannetje
vermiljoenrood en het wijfje bruinachtig groen is. Indien nu
schitterende kleuren uiterst gevaarlijk voor de vogels waren
geweest, terwijl zij op hun open nesten zaten, [161]zouden de
mannetjes in deze gevallen zeer hebben geleden. Het zou echter
voor het mannetje zoo belangrijk kunnen zijn om schitterend
gekleurd te wezen, dat dit meer dan opwoog tegen een weinig
daardoor veroorzaakt grooter gevaar.
De heer Wallace geeft toe, dat bij de Koningskraaien (Dicrurus),

Wielewalen (Oriolus) en de Aardlijsters (Pittidae) (2) de wijfjes
opzichtig zijn gekleurd en toch open nesten bouwen; maar hij wijst er
op, dat de vogels van de eerste groep zeer strijdlustig zijn en zich
zouden kunnen verdedigen; dat die van de tweede groep de uiterste
zorg aanwenden om hun open nesten te verbergen (doch dit houdt
niet altijd steek 12), en dat bij de vogels van de derde groep de wijfjes
voornamelijk op de ondervlakte van het lichaam levendig zijn
gekleurd. Behalve deze gevallen maakt de geheele groote Familie
der Duiven die somtijds levendig en bijna altijd opzichtig zijn
gekleurd, en van welke het algemeen bekend is, dat zij van de
aanvallen der roofvogels hebben te lijden, een ernstige uitzondering
op den regel, want duiven bouwen altijd open en blootgestelde
nesten. In een andere groote Familie, die der Kolibri’s, bouwen al de
soorten open nesten; toch zijn bij eenige van de prachtigste soorten
de seksen op de zelfde wijze gekleurd; en bij het meerendeel zijn de
wijfjes, hoewel minder schitterend dan de mannetjes, toch zeer
levendig gekleurd. Men kan ook niet volhouden, dat alle vrouwelijke
kolibri’s die levendig zijn gekleurd, aan de ontdekking ontsnappen,
omdat zij groen zijn; want sommige prijken op de bovenvlakte van
hun lichaam met roode, blauwe en andere kleuren. 13
Wat de vogels betreft, die in gaten bouwen of koepelvormige nesten

bouwen, zoo worden hierdoor, gelijk de heer Wallace opmerkt, nog
andere voordeelen dan verberging verkregen, zooals beschutting
voor den regen, grootere warmte en in heete landen bescherming
voor stralen der zon 14, zoodat het geen geldige tegenwerping tegen
deze meening is, dat vele [162]vogels, bij welke beide seksen donker
zijn gekleurd, verborgen nesten bouwen. 15 De vrouwelijke
Neushorenvogels (Buceros), bij voorbeeld, van Indië en Afrika
worden gedurende den nestbouw (3) met bijzondere zorg
beschermd; want het mannetje metselt het gat dicht, waarin het
wijfje op de eieren zit, en laat alleen een kleine opening over, door
welke hij haar voedt; zij wordt dus gedurende den geheelen broeitijd
in een enge gevangenis opgesloten 16; en toch zijn de vrouwelijke
Neushorenvogels niet opzichtiger gekleurd dan vele andere vogels
van gelijke grootte, die open nesten bouwen. Het is een ernstige
tegenwerping tegen de meening van den heer Wallace, gelijk hij ook
toegeeft, dat in eenige weinige groepen de mannetjes schitterend en
de wijfjes donker zijn gekleurd en deze laatste toch de eieren in
koepelvormige nesten uitbroeien. Dit is het geval bij de Grallinae van
Australië (4), de Prachtzangers (Maluridae) van het zelfde land, de
Zon-vogels (Nectariniae) en met verscheidene van de Australische
Honigzuigers of Meliphagidae. 17
Indien wij de vogels van Engeland beschouwen, zullen wij zien, dat
er geen nauw en algemeen verband bestaat tusschen de kleuren
van het wijfje en den aard van het door haar gebouwde nest.
Omtrent veertig van onze Britsche vogels (die van aanzienlijke

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Animals and the Shaping of Modern

Medicine: One Health and its Histories

Shaping the Geography of Empire: Man and Nature in

Conversion Narratives in Early Modern England: Tales of

Medicine, Magic and Art in Early Modern Norway 1st ed.

The Struggle for Health: Medicine and the politics of

English Grammar All-in-One For Dummies 1st Edition

The Dog: Its Behavior, Nutrition, and Health 2nd

Pandemic Police Power, Public Health and the Abolition

Oxford Case Histories in Obstetric Medicine Charlotte

ABIGAIL WOODS, MICHAEL BRESALIER,

More information about this series at

Animals and the

Michael Bresalier Rachel Mason Dentinger

Medicine and Biomedical Sciences in Modern History

Library of Congress Control Number: 2017948270

Printed on acid-free paper

This Palgrave Macmillan imprint is published by Springer Nature

1Programme Grant reference 092719/Z/10/A.

findings in the course of the research programme. Abigail Woods would

London, UK Abigail Woods

1 Introduction: Centring Animals Within Medical History 1

2 Doctors in the Zoo: Connecting Human and Animal

3 From Coordinated Campaigns to Watertight

4 From Healthy Cows to Healthy Humans: Integrated

5 The Parasitological Pursuit: Crossing Species and

6 Humans, Other Animals and ‘One Health’ in the Early

6.1.3 The Swiss Tropical and Public Health Institute 207

Appendix: Annotated Bibliography of Animals

Fig. 6.1 The One Health umbrella 199

Table 3.1 British research into diseased sheep, 1880–1901 78

Introduction: Centring Animals

In a recent handbook on the history of medicine, authors Robert Kirk

to a limited but important extent, writing about human history is always—

However, Benson acknowledges the difference between scholarship that

Woods (2016, 2017a, 2017b), Kirk (2017).

© The Author(s) 2018 1

Expanding beyond much-studied laboratory contexts to explore the

Control and Prevention, One Health Initiative.

that are important to the writing of animal histories, we present them

1.1 Why Animals?

actors in the drama of human history. Their stated intention is to bring

9 Ritvo (2002), Fudge (2002), Kean (2012), Sivasundaram (2015).

for making this observation.

13 For example: Bekoff (2002), Grandin and Johnson (2005).

of environments and ecosystems, environmental historians have explored

14 McNeill (2000), Nash (2005), Cronon (1990).

18 Taylor and Twine (2014), Institute for Critical Animal Studies.

and the ‘resistance, compliance and compromise’ that variously charac-

1.2 Writing Animal Histories

20 Pooley-Ebert (2015) p. 165.

26 Philo and Wilbert (2000), Eitler (2014), Pooley-Ebert (2015).

27 Callon and Law (1986), Latour (1988, 2005).

of relations (or networks) in which all are embedded. The configuration

28 Pearson (2013) pp. 128–45, Eitler (2014).

32 Pettit et al. (2015).

1.3 Animals in Medical History

35 Kirk and Worboys (2011).

veterinary history, which is ostensibly focused on the health of animals as

1.3 Animals in Medical History