2008 colofon n 4:2005 COLOFON 3/4

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PARASSITOLOGIA
A publication of “Sapienza” University of Rome
Official Journal of the Italian Society of Parasitology
EDITOR-IN-CHIEF
M. Coluzzi
Volume 50, No. 3-4
December 2008
ASSOCIATE/CORRESPONDING EDITORS
General Parasitology
Veterinary Parasitology
Medical Parasitology
Molecular Parasitology
Sanitary Entomology
L. Sacchi
G. Cringoli/D. Otranto
F. Bruschi/E. Pozio
C. Bandi
A. della Torre
EDITORIAL BOARD
The Council (2008-2012) of the Italian Society of
Parasitology: F. Bruschi (Vice-Presidente), S. Cacciò (Tesoriere), G. Cringoli (Membro), F. Esposito
(Membro), E. Ferroglio (Membro), A. Frangipane
di Regalbono (Segretario Generale), M. Pietrobelli
(Presidente), G. Poglayen (Membro)
ADVISORY BOARD
A. Aeschlimann, P. Ambroise-Thomas, H. Babiker,
V. Baimai, D.J. Bradley, R. Carter, A. Chabaud,
C. Combes, C. Curtis, J. de Zulueta, K. Dietz,
J.P. Dubey, T.H. Freyvogel, B.M. Greenwood,
C. Louis, K. Marsh, S.A. Nadler, R.S. Nussenzweig,
I. Paperna, J.M.E. Ribeiro, J.A. Rioux, D.
Rollinson, R. Roncalli, M.W. Service, J.D. Smyth,
Y.T. Touré, J. Vercruysse, D. Wakelin, G.B. White
EDITORIAL OFFICE
Dipartimento di Scienze di Sanità Pubblica
Sezione di Parassitologia “Ettore Biocca”
Università “Sapienza” di Roma
Piazzale Aldo Moro 5, I-00185 Roma, Italy
Tel ++39 06 4455780
Fax ++39 06 49914653
e-mail: mario.coluzzi@uniroma1.it
CONTENTS
INSECTS AND ILLNESSES: CONTRIBUTIONS
TO THE HISTORY OF MEDICAL ENTOMOLOGY
A Conference in three Sessions:
London, April 2005; Paris, April 2006; Rome, October 2007
Editors’ Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
1. Construction of the reference insects collections
B. BACCETTI - History of the early dipteran systematics in Italy:
from Lyncei to Battista Grassi . . . . . . . . . . . . . . . . . . . . . . . 167
Y. CAMBEFORT - Knowledge of Diptera in France from the beginning to the early twentieth century . . . . . . . . . . . . . . . . . . . . 173
M. ROMERO SÁ - Scientific collections, Tropical Medicine and the
development of Entomology in Brazil: the contribution of Instituto Oswaldo Cruz . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
2. Portraits
E. CAPANNA - Battista Grassi entomologist and the Roman School
of Malariology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
M.A. OSBORNE - Raphaël Blanchard, Parasitology, and the positioning of Medical Entomology in Paris . . . . . . . . . . . . . . . . 213
J.-P. DEDET - The Sergent brothers and the antimalarial campaigns
in Algeria (1902-1948) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
F. DELAPORTE - The discovery of the vector of Robles disease . . . 227
PUBLISHER
Lombardo Editore, Divisione Periodici
Production and Subscription Offices:
Via Centrale 89 (Lama),
I-06013 San Giustino (PG), Italy
Tel ++39 075 8583860
Fax ++39 075 8610415
e-mail: infolombardo@lombardoeditore.it
J.L. BENCHIMOL - Medical and Agricultural Entomology in Brazil:
a historical approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
3. Insects variation and adaptation to environment
R. HOUIN - Culicoides and the Tartar Steppe: Il Deserto dei Tartari. Culicoides and the spread of blue tongue virus . . . . . . 249
(continued)
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Contents
II
P
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A
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S
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Founded in 1959 by
E. Biocca, A. Corradetti and O. Starkoff
A. OPINEL - Reconstructing an epistemological itinerary: environmental theories of variation in Roubaud’s experiments on
Glossina flies and Anopheles, 1900-1938 . . . . . . . . . . . . . . . 255
G. GACHELIN , A. OPINEL - Theories of Genetics and Evolution and the
development of Medical Entomology in France (1900-1939) 267
4. Social and economical perspectives on Medical Entomology
T. GILES-VERNICK - Entomology in Translation: Interpreting French
medical entomological knowledge in colonial Mali . . . . . . . . 281
D. GILFOYLE - Science and popular participation in the investigation of heartwater in South Africa, c. 1870-1950 . . . . . . . . 291
K. BROWN - Veterinary Entomology, colonial science and the challenge of tick-borne diseases in South Africa during the late
nineteenth and early twentieth centuries . . . . . . . . . . . . . . . . 305
J.F.M. CLARK - Sowing the seeds of Economic Entomology: houseflies and the emergence of Medical Entomology in Britain . . . 321
List of participants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329
FRONTESPIZIO 1 2008:FRONTESPIZIO 3/4 2005
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INSECTS AND ILLNESSES: CONTRIBUTIONS
TO THE HISTORY OF MEDICAL ENTOMOLOGY
A Conference in three Sessions:
Wellcome Trust Centre for the History of Medicine at University College of London, April 2005
Institut Pasteur, Paris, April 2006
Accademia dei Lincei, Rome, October 2007
edited by
Mario Coluzzi
Università “Sapienza” and Accademia dei Lincei, Rome
Gabriel Gachelin
Rehseis, CNRS/Université, Paris
Anne Hardy
The Wellcome Trust Centre for History of Medicine at University College of London
Annick Opinel
Centre de recherches historiques, Institut Pasteur, Paris
LOMBARDO EDITORE
FRONTESPIZIO 1 2008:FRONTESPIZIO 3/4 2005
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DEDICATION
This double issue of Parassitologia is dedicated to Teresa Ariaudo
who suddenly was taken away (5 December 1957 - 17 February 2008)
while was working at the editing of the present fiftieth anniversary volume of
our journal, for which she acted as secretarial assistant. All those who had
working relationship with her have appreciated her kindness availability,
precision, cultural and ethical qualities.
1 GUEST EDITORS:1 preliminari 3/4 2005
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Parassitologia 50 : 157-163, 2008
Editors’ Introduction
The birth of medical entomology, even if not defined with this formal name, is generally put at the
beginning of the big discoveries of transmission of diseases by insects and ticks, i.e. the discovery
by Manson of the transmission by mosquitoes of bancroftian filariasis dated 18771. However we
should not forget the early studies on Acarus scabiei which contributed to overcome the
Hippocratic medical tradition by establishing the parasitological nature of scabies infection. Much
time before the Pasteurian revolution, the hypothesis of the “contagium vivum” had a possible
experimental verification in the study of pathologies from arthropods ectoparasites and it was in
particular the parasitic nature of scabies to constitute the subject for long confrontation between
supporters and adversaries of the “patologia animata”. In 1687 a letter by Cosimo Buonomo to his
master Francesco Redi is published in which it is given the description of the parasite clearly indicated as the etiological agent of the dermatosis. This letter is based on observations of the author
and of Djacinto Cestoni, who were respectively physician and pharmacist in Livorno. In 1746
Linneo confirmed the parasitological findings, listing among the species of the fauna suedica
Acarus exulcerans, later named Acarus scabiei, of which is said “habitat in scabie ferine cuius
causa est”.
These first essays in medical entomology are followed by the numerous studies on insects which
started with the discovery of the compound microscope by Galileo Galilei which provided the
basic of systematic entomology on which the specialized science of medical entomology was built
as is illustrated in this issue by Benchimol, Baccetti and others.
Raphaël Blanchard, the distinguished Professor of Parasitology and Medical Zoology at the Faculty of
Medicine of Paris from 1883 to 1919, was the first officially to define Medical Entomology
(Entomologie médicale) in his general address to participants at the First International Congress of
Entomology, held in Brussels in 1910. For Blanchard, Medical Entomology was that part of
Entomology dealing with the insects responsible for arthropod-borne diseases, particularly parasitic
diseases2. This definition may have been in circulation before 1910, but Blanchard had not used the
expression Entomologie médicale in his 1890 reference book on Medical Zoology. Instead, he then
described insects as mere nuisances, and conceded only few pages of his volume to the biting
diptera3. Over the course of twenty years, however, it had become necessary to qualify a subfield of
Entomology, generating a compound name that was immediately understood and accepted by physicians and veterinarians. It appears that the rapidly acquired diverse knowledge about insects of medical interest and the distinctive features of Insect Biology and Taxonomy in transmitting some diseases, somehow led to the “self-organization” of a specific field of research and teaching. Since then,
the expression “Medical Entomology” has been used continuously, and has been widely accepted.
The medical importance of insects was only vaguely recognised from 1880, after insects were identified as vectors of yellow fever and filariasis, more so during the last five years of the nineteenth
century. Its real significance, however, increased only after the medical and scientific communities
generally accepted insects as vectors around 1900, and once these insects had been recognised as
transmitters of malaria, yellow fever, plague and sleeping sickness, and contributors to the life
cycles of parasites. The last quarter of the nineteenth century was critical, for it was then that the
foundational knowledge of Medical Entomology was acquired. The production of this knowledge
was itself embedded in a plurality of other developments during this period, including the birth of
the notion of “Tropical Medicine”, the systematisation of Microbiology and Parasitology, and the
M.W. Service (1978). Patrick Manson and the story of bancroftian filariasis. Royal Society of Tropical Medicine and Hygiene in Medical Entomology Centenary, 11-14.
2 A. Opinel (2008). The emergence of French medical entomology: the respective influence of universities, Institut Pasteur and army physicians (ca 1890 to 1938). Med Hist 52(3): 387-405.
3 R. Blanchard (1890). Traité de zoologie médicale, J.B. Baillière et fils, Paris.
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importance granted to Evolution, Genetics and Ecosystems (biocenoses). New or more aggressive
commercial and colonial policies were also significant. The general acceptance of the expression
“Medical Entomology” before World War I thus reflects the prominent role played by insects in the
economic life of tropical (and some other) countries, as expressed at the turn of the twentieth century by the organization of specific instruction in tropical medicine in several European countries.
Numerous historical studies exist of Tropical Medicine, its teaching 4, and links with empire, trade
and local public health 5. There are historical studies of several arthropod-borne viruses, and the
bibliography on malaria fills library shelves. Similarly, there are countless biographies of the best
known historical actors connected with these diseases. However, with such exceptions as Adolpho
Lutz, these actors are better known as parasitologists than as the entomologists they had become.
Historians, as well as scientists, consider arthropod-borne diseases other than malaria, Chagas disease and yellow fever, to be neglected diseases. As a whole, scholars have addressed Medical
Entomology implicitly rather than analysing the field outright. Research and practices in entomology as applied to medicine thus appear to have received less attention from a historical perspective
than the diseases that insects transmit. This historical neglect of Medical Entomology prompted us
to gather contributions that illustrate the strategies used, and the challenges faced, by entomologists working on the vectors of various diseases. That approach defined an epistemological strategy of Medical and Veterinarian Entomology which differed from standard microbiological studies.
This strategy was closer to those used in biological studies of natural, complex ecosystems. In turn,
some scientific problems of basic biology, namely the origin of biological variation of organisms,
became key issues in medicine, for they were closely linked to the designing efficient and economical forms of disease prophylaxis. The present issue of Parassitologia is thus not intended to be a
history of Medical Entomology, nor does pretend to offer an exhaustive analysis, since many vectors and historical actors are not addressed. Rather, this issue seeks to explore the complexity of
the technical and biological problems that medical entomologists faced in their attempts to elucidate and control the relations between a parasite or microbe, its vector and its vertebrate victims.
Most contributors to this issue have participated in at least two of the three workshops organized
on this theme, the first at the Wellcome Trust Centre for the History of Medicine at University
College of London (April 2005), the second at the Institut Pasteur in Paris (April 2006) and the
third at the Accademia dei Lincei in Rome (October 2007). Each paper synthesizes several successive presentations, benefiting from these several opportunities to discuss various approaches to a
given perspective – medical, veterinarian, economic, fundamental – on entomology.
Medical Entomology takes its legitimacy from medicine, but is equally rooted in entomological
knowledge. Even physicians must provide precise names of the insects carrying the germs of disease that they identify and cure, and they undertake this task within the framework of the Western
taxonomy. The elusive meaning of the African denominations of malaria and its vector, described
by Tamara Giles-Vernick in the present volume, well illustrates the fact that denominations in
other cultures reflect the altogether different properties attributed to the disease and to the vectors
in those cultures. Indeed, such denominations are part of a distinctive cosmogony or culture,
which attribute specific meanings to particular societies’ immediate environments. A similar problem is associated with sickle cell anemia in Africa, and is part of the difficulties routinely encountered by Western-trained physicians in their attempts to correlate their own interpretations of
symptoms with those used by local populations. In this volume, John Clark discusses the importance granted to the propagation of infection by house flies, considered a domestic enemy, and
4 The contribution of L. Wilkinson on the London School of Hygiene, presented during the first workshop in
London, is not included. Readers are invited to refer: L. Wilkinson and A. Hardy (1999). Prevention and Cure:
The London School of Hygiene & Tropical Medicine, a Twentieth-Century Quest for Global Public Health, Kegan
Paul, London.
5 The manner arthropod borne diseases had been identified and fought in Brazil, including a discussion on the
role of triatoma and phlebotomes, has been earlier discussed in: A. Opinel and G. Gachelin, Eds (2005). Parasitic
diseases in Brazil: the construction of Parasitology, XIXth and XXth centuries. Parassitologia 47: 255-395.
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suggests that behind the hygienic rationality of house fly eradication lies an array of less rational
attitudes. Such attitudes can be seen in the metaphoric use of images of parasites and insects in
post World War I pamphlets and posters. Medical Entomology is fundamentally a Western-based
science, and entomology relies on taxonomy, which obeys rules established during the nineteenth
century, according to Yves Cambefort. The criteria can change, but the principles of a taxonomy
based on binary characters do not. That said, collections were the basis on which insect taxonomy
developed in Europe during the nineteenth century, as both Yves Cambefort and Baccio Baccetti
remind us. All this work of describing and storing insect types resulted in an extraordinary knowledge, but museum and library specialists kept it so well concealed in their own institutions that it
was useless to physicians. These physicians had to struggle alone to become medical entomologists
and to publish their own reference atlases for insect identification.
An unexpected conclusion is raised in papers by Baccetti, Cambefort, Jaime Benchimol and Magali
Romero Sà. Despite a permanent exchange of data, samples and even complete collections of insects
between different countries and researchers, the policies of specimen collection and the organizing
of reference collections of insects of medical interest after 1900 displayed national features. As soon
as “mosquitoes” (not used here as a correct entomological denomination!) were shown to carry parasites and microbes, Theobald, on behalf of the Natural History Museum in London, launched in
1901 a world wide survey of diptera and other insects of medical interest on the basis of an economic and imperial rationale. Howard and the United States Army launched a similar survey of insects
present in Central and South America which they considered to be their natural sphere of influence
under the Monroe Doctrine. Benchimol describes the systematic survey of insects of medical interest
present in the Brazilian territory, which was carried out by Lutz, Neiva, Goeldi and colleagues from
the beginning of the twentieth century, and which produced reference collections, most of which are
kept at Institute Oswaldo Cruz. Public health was the leading motive for this survey and collection.
Similar conclusions, not discussed in the present issue, can be made concerning Germany and
Belgium. Italy, as both Baccetti and Ernesto Capanna note, had been engaged for millenia in a struggle against malaria, which eventually resulted in a national policy towards the disease, a well-developed infrastructure to control its transmission (see for example the laws on bonifica), and the
administration of quinine. As soon as the role of Anophelines was known, the fight against them
was incorporated into these measures. As Capanna reveals, Malariology was part of Italian scientific
culture, and it reflected significant national pride, conferring on Grassi and his colleagues a tremendous importance. By contrast, a national policy was absent in France, where the Museum National
d’Histoire Naturelle failed to retain Bigot’s reference collection of diptera (moved to England in
1893), despite a rapidly-expanding colonial empire. The lack of a French national policy might have
been due to the fact that the French showed little interest in their colonies, except for the settler
colony of Algeria, where malaria was of singular importance. That absence of a coordinated national
policy was to some extent compensated for by individual institutional initiatives. French Medical
Entomology developed in the field through missions and reached international standards very quickly through the efforts of scientists and physicians working in a few institutions, notably Blanchard at
the Faculté de Médecine de Paris (see the papers by Osborne 2008, Opinel 2008, present issue), the
Institut Pasteur (Opinel 2008) and the Pasteur institutes overseas, particularly the Institut Pasteur in
Algiers as described by Dedet but also those in Madagascar, Indochina and the West Indies.
Moreover an important contribution to French Medical Entomology came from ORSTOM (Office
de la recherche scientifique et technique Outre-mer) organization that now continues thanks to the
efforts of the new Institute of Research for Development. In spite of their focus on ex-colonial territories, this organization has given to Medical Entomology in France the needed international standards and the colleagues of the IRD are undoubtedly competitive with the best centres of Medical
Entomology world wide. It must also to be said that important chairs of Parasitology in France particularly the chair of Strasbourg and that of Montpellier have taken in great priority the Medical
Entomology programme contributing important work in this field. But a most important contribution was the one of Sergent brothers on malaria control in Algeria. Their work well summarized by
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The illustrations on male and female adults of Aedes aegypti, vector of yellow fever, is reported from plate 1 of the book of
Emilio Augusto Goeldi. Unrivaled precision of the drawings shows well the level reached by these South American colleagues already at the beginning of the last century (Goeldi EA, 1905, Memorias do Museu Goeldi “Os Mosquitos no Pará”).
Dedet in this issue contains all the main topics of vector control and provides the first proof of
malaria eradication as a feasible intervention in temperate zone ecosystem.
Standard research practices were soon established within Medical Entomology, permitting the easier identification of a vector whenever a virus or parasite was suspected as the causal agent of a disease. In this respect, the identification of the vector of oncocerchiasis between 1914 and 1916 by
Rodolfo Robles, described by François Delaporte (2008), is an exemplary case of such research in
the field.
Medical Entomology includes Veterinary Entomology. Animals can develop diseases similar to
those of human beings, diseases of their own, or remain healthy carriers of disease. Arthropod
borne microbes often cause animal diseases, with serious economic consequences where they
affect cattle. Papers by Karen Brown and Daniel Gilfoyle deal with the development of Veterinary
Entomology in South Africa. Both analyse the role of ticks in the transmission of different diseases. The manner of identifying the tick species transmitting a particular microbe is by no means
different from the process of identifying a human disease vector. The point is that local prevalence
of a bacterial or parasitic disease, transmitted by ticks to commercially valuable cattle, required
the application of prophylaxis and therapies by methods that could never be used among human
populations. Ticks were part of the South African bush ecosystem, and the only way to decrease
the propagation of tick borne diseases was to kill ticks just as they attached themselves to the animals, and simultaneously to clean up lesions through which diseases were transmitted. This control strategy was achieved through the development of pediluves and of trenches filled with arsenic
derivatives, which served as efficient external pesticides. Both Brown and Gilfoyle discuss the
rationale, procedures, and results, as well as local populations’ resistance to these measures. The
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most striking feature of these case studies are the mass procedures used and the reliance on chemistry, rather than on prophylactic measures to tackle the vector or its environment.
The essays in this volume suggest a further conclusion: that medical entomologists and physicians
had to account for the ways in which complex ecosystems sustained disease transmission. Thus,
whenever a disease became endemic in a particular ecosystem, the physician had to intervene in
the infected ecosystem to interrupt the cycle of transmission. The reliance on chemicals, despite
their frequent toxicity and the need for repeated treatments, could be a logical and efficient manner of interrupting the microbe’s biological cycle and progressively reducing transmission frequency. A less dramatic alternative was to identify those features of the ecosystem which might be
influenced to induce a decline in the propagation of vector and microbe. The drainage of marshes
and pestilent areas was practiced in Italy from Etruscan times; the drainage of the Pontine marshes
and the Po valley were the goals of Mussolini’s Grande bonifica. Public works to drain and dry
infected areas, so as to reduce those spaces in which diptera could develop, were common practice
throughout the world. But these procedures were costly. Careful descriptions of insects, human
beings, and parasites in their environments offered different approaches. It was after preliminary
studies of Glossina palpalis behaviour, and principally the identification of the wooded areas
where these insects preferred to live, that Brumpt in 1903 proposed the establishment of villages
at some distance from water flows and the deforestation of river embankments. The experiments
carried out in the Mitidja valley by Edmond and Etienne Sergent between the two World Wars,
and described in this volume by Jean-Pierre Dedet, drew their inspiration from Grassi’s research.
These experiments studied the local Anopheles’ biological cycle and used the alternation of water
flows to prevent the larvae from maturating. The careful environmental descriptions, carried out
by researchers who borrowed Rockefeller Foundation strategies, sought to identify precisely the
areas to be treated, and to define which prophylactic measures (biological, drainage or chemical
spraying) were most appropriate. Applied in Corsica by Brumpt and in Italian marshes by
Hackett 6, the environmental analysis included all local parameters then thought to influence the
infectivity of Anopheles. These studies were carried out at a time when debates about the zooprophylaxis hypothesis raged and when the taxonomic status of Anopheline species were being
defined. Emile Roubaud, whose studies are analysed by Annick Opinel in the present issue, contributed extensive descriptions of the environmental influences on Glossina flies before World
War I; after the war, he developed analogous assessments of Anopheles. Roubaud developed several hypotheses concerning the direct environmental influences on a vector’s capacity to transmit disease, including the nature of its food and of climatic parameters, the emergence of the specific biological features (feeding and biting behaviour that led to preferences for feeding on cattle responsible for zooprophylaxis, housing and mating behaviours, etc.). Would it possible to “educate”
insects properly, Roubaud believed, transmission between insects and human beings, and vice
versa, could be controlled. During the first third of the twentieth century, this possibility led him
to carry out experiments that sought to modify the behaviours of dangerous insects in the laboratory, and to “educate” them to become less dangerous, if not innocuous, to human beings.
Roubaud was a follower of Lamarkian theories and as such he had a negative influence on French
Medical Entomology between the two Wars. His negative influence is reflected in the work of
some of his students as it was the case of Max Holstein. To this medical entomologist a staff in
WHO was given by Professor Pampana at the time director of malaria eradication division, a wonderful opportunity to work with Professor Frizzi on the Anopheles gambiae complex. Frizzi and
Holstein were the first to see some of the chromosomal variations in An. gambiae and in An. arabiensis (1956), at that time not yet described by George Davidson as species A and B. The influence of Holstein was very negative because he ignored the speciation phenomena and he described
different species under the Linnean name An. gambiae. In a later publication (1959) he also
attempted to put in synonymy An. melas with An. gambiae, the only two species of the complex
A. Opinel and G. Gachelin (2004). The Rockefeller Foundation and the prevention of malaria in Corsica
(1925-1931): the support to the French parasitologist Emile Brumpt. Parassitologia 46: 287-302.
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well distinct ecologically being An. melas a salt water breeder which will be also recognized on
morphological ground. What Max Holstein was unable to understand because of the imprinting of
Roubaud was something which appeared clear to a non-entomologist, medical trained men, like
Lewis Hackett (1937) who in his book “Malaria in Europe” wrote “It will help us to understand
the diversified behaviour of the varieties of maculipennis if we can forget their extraordinary physical similarity and treat them as different species”.
Whatever resulted from the debates about the effects of zooprophylaxis and insect education on sleeping sickness and malaria transmission, nearly all medical entomologists had to confront an idea about
environment that was novel to them, and to most biologists. Parasites and insects were adapted to
local conditions, and their environmental adaptations were responsible for the frequency of diseases.
The nature of the adaptation processes underpinning the emergence of biological species was extensively discussed until World War II. The genetic and evolutionary context in which these researches
and discussions were pursued is described in a contribution by Gabriel Gachelin and Annick Opinel
in this present issue. These insights replaced older dichotomous understandings of acquired and
selected characteristics, and contributed to the over stressing of the notion of biological species in
most cases defined only by chromosomal polymorphisms (inversions). Despite the resolution of the
Anopheles maculipennis complex in 1935, the interpretation of the data collected in the field
remained difficult until Mario Coluzzi, with his researches on the An. gambiae complex provided
genetic evidence linking the observed selected adaptation to a given, precise, environment and to
species, sometimes only incipient but marked by genetic rearrangements of Anopheline chromosomes.
The role of genetic variations in the microbe and in its vector(s) is confirmed in René Houin’s descriptions of the various stages in the progression of blue tongue disease from Africa to Great Britain and
continental Europe. This journey involved the selection of an aggressive variant of the virus, followed
by its passage into a new vector adapted to temperate climates. The example of blue tongue disease
provides a good model for the possible spread of arthropod borne human diseases from their usual
areas of distribution into other climatic zones irrespectively of global warming.
The contribution of Mario Coluzzi was particularly important in terms of genetics and speciation.
He starts with a modern definition of species that he is intended as a group of organisms actually
or potentially interbreeding which is separated by other similar groups by intrinsic mechanisms of
reproductive isolation. So these species are seen first of all as a panmictic unit constituted by individuals reproductively compatible, which are expressed by, and contributing to the variability of
the same gene pool. This definition shows the species as a dynamic, not static, category. The speciation as described by Coluzzi in the An. gambiae complex is always a phenomenon associated to a
genetic polymorphism (chromosomal inversions) which constitute a series of adaptive options
build by the species in its past experience while testing different environments. This process called
of “ecotypification” by which the various adaptive options are represented by cytodemes and as
such would be ready to expand the normal range of the species in its attempt to adapt permanently
to a marginal environment. These adaptations are constructed thanks to the inversions or any
other mechanism of cross-over suppression (Coluzzi, in press).
A concept of species and of cryptic species which is closely related to recent cases of speciation
and sometimes to incipient species was from the beginning in the background of malariological
entomology particularly concerning the problem of anophelism without malaria. This problem was
a specially puzzling in central Italy were An. labranchiae was associated with its sibling An.
atroparvus as well as with other taxa of maculipennis complex. Now we know that An. labranchiae is the most termophilic species of the complex being originary of North Africa and only secondarily expanded its range of distribution occupying northern cooler areas. Expanding to the north
An. labranchiae had no difficulty when reaching Sardinia since it was the only member of the
maculipennis complex on this island. More problematic was for the species to expand along the
Italian peninsula where, in fact, An. labranchiae was involved in some niche partitioning having a
costal distribution in agreement with its termophily while the other species of maculipennis complex had mostly an inland distribution.
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An. labranchiae was also more associated with men and his environment, more anthropophilic
and more important malariologically particularly in the case of transmission of Plasmodium falciparum since it was the only species of the maculipennis complex without rigid diapause but able
to overcome the winter season with only gonotrophic dissociation. This incidentally may explain
the dramatic impact of indoor sprayed insecticides, DDT, which forced the Anopheles outdoor
with its irritant effect making the species unable to survive during the winter time particularly in
the northern border of its distribution range. About this border it has been in central Italy and
despite of minor fluctuations corresponding to the northern border of Grosseto province. There
was something which avoided the expansion of An. labranchiae to the north, a factor undoubtedly
related to the phenomenon of competitive exclusion by one or more species of the maculipennis
complex presumably the sibling An. atroparvus having a major role. We can really understand how
difficult it was this situation to interpret malariologically without having the background knowledge pointed out above. Why Grosseto province was the last to have serious malaria problems in
spite of the fact that what was regarded as apparently identical species of Anopheles were well
present also in central and northern Tuscany? A very confusing story for our old masters Celli and
Grassi, the real enigma known as anophelism without malaria.
Taken as a whole, the diverse contributions to the present issue of Parassitologia point to the conclusion that Medical Entomology operates within a context in which all biological partners are in constant interaction, but are also dependent on the physical features of a distinctive environment, including its fauna and flora, and habitat. The studies of Medical Entomology cannot be dissociated from
the study of ecosystems. Any medical entomologist will immediately identify a breeding place for
Anopheles maculipennis, or for Glossina palpalis or others, simply by looking at a landscape or even
a picture of one. Modern biologists searching for mechanisms common to all members of a genus,
and susceptible for targetting by drugs, need to understand the diversity of the situations encountered by medical entomologists at the beginning of the twentieth century, and of the remarkable
adaptive properties of the insects and parasites discussed in the present issue of Parassitologia.
MARIO COLUZZI, Università “Sapienza” and Accademia dei Lincei, Rome
GABRIEL GACHELIN, Rehseis, CNRS/Université, Paris
ANNE HARDY, The Wellcome Trust Centre for History of Medicine at University College of London
ANNICK OPINEL, Centre de recherches historiques, Institut Pasteur, Paris
1 OCCHIELLO:1 OCCHIELLO
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26-08-2009
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1
Construction of the
reference insects collections
26-08-2009
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Pagina 165
History of the early dipteran systematics in Italy:
from Lyncei to Battista Grassi
B. Baccetti
Emeritus Professor, University of Siena, Italy.
Abstract. This presentation starts with Galileo’s discovery of the microscope and the first Lyncei. Giovanni Heckius and Francesco Stelluti demonstrated different kinds of mosquitoes. Later, in Florence, the
Academy of Cimento solved the problem of mosquito reproduction with the discoveries of Francesco
Redi, Pietro Paolo da Sangallo, Giuseppe Del Papa and Giovanni Maria Lancisi in the 18th century. In
19th century Eugenio Ficalbi reviewed the Italian Culicids. Once Battista Grassi solved the cycle of
Anopheles and Plasmodia, further researches followed by Golgi, Celli, Marchiafava, Bastianelli and Bignami, as well as by Roland Ross.
Key words: dipteran systematics, Italy, malariology, history.
The first description of mosquitoes was obtained by
the early microscopists, and in particular by the
founders of the Academy of Lyncei, immediately
after Giovanni Heckius returned from his visit to
Francesco Stelluti (1577-1653) during which
showed him some beautiful pictures of insects.
Insects including mosquitoes were subsequently
studied under the Galilean microscope. Stelluti participated in the final preparation of the material presented at the first Lyncean meetings, before 1616.
But the Lyncei had a short life, and was replaced by
the “Medici Accademia del Cimento” in Florence.
One evening in June 1679, a group of Florentine
philosophy students were walking by the foundation
of the Basilica di Santa Maria del Fiore, amicably discussing scientific subjects, when some mosquitoes,
which were buzzing around them, attracted their
attention. The youngest of these scholars was Pietro
Paolo da Sangallo, and all were students and friends
of Francesco Redi (born in Arezzo, 1626 and died in
Pisa, 1697) very well-known in the Florence at that
time, a member of the Crusca and Cimento Academies, professor at the University of Pisa, a chief
physician of the Grand Dukes de’ Medici, first of Ferdinand II and then of Cosimo III1.
A year after the closure of the Cimento Academy,
Redi managed to publish his book Experiments in
the generation of Insects (Florence, 1668, “all’Insegna della Stella”) in the form of a letter to Carlo
Dati, defining himself on the title page as a member
of the Crusca Academy, without mentioning the
Cimento. This work was reprinted many times. Five
editions appeared in Florence by 1688, and a Latin
Correspondence: Baccio Baccetti, Section of Biology, University of Siena, S. Maria alle Scotte Hospital, Lotto 1, Piano 1S,
53100 Siena, Italy, e-mail: baccetti@unisi.it
1 Ferdinand II was also the founder of the Cimento Academy in 1657 and, with his brother Leopoldo, he was the patron
and one of the most active members during the life of the
Academy up until 1667, when it was closed, as Leopoldo was
elected cardinal and consequently called to Rome.
translation was published in Amsterdam in 1671.
With these experiments, Redi was the first to
demonstrate the erroneous nature of belief in spontaneous generation. This brought controversy, since
it ran counter to the official view of the Church,
supported by the Jesuits Atanasio Kircher and Filippo Buonanni. Moreover it seems that the Vatican,
which had for some time been annoyed by Redi’s
innovative ideas, took caution by creating Leopoldo
de’ Medici a cardinal and summoning him to Rome,
so silencing the Cimento Academy.
At the time when the little group of students discussed mosquitoes, Redi was their acknowledged
master and spiritual father, he had already published
the Experiments, and he had many well-known students in Florence who dedicated their studies to
him, including Lorenzo Bellini, Giuseppe del Papa
(who succeeded him as chief physician to the Grand
Duke and professor at Pisa), Giovanni Caldesi, an
academic who studied tortoises, Pietro Paolo da
Sangallo, who studied mosquitoes, and finally the
circle in Livorno which consisted of Giovanni Cosimo Bonomo and Diacinto Cestoni, who, togheter
with Redi, demonstrated the parasitic origin of scab
(Bonomo, 1687, pp. 1-13).
Following the closure of the Cimento Academy, it
was due to this group of scholars that the collaborative spirit which was typical of that Academy survived in Florence. Da Sangallo gave proof of that
spirit, writing to Redi to explain how he had started his studies of mosquitoes, as he had confided to
his friends that famous evening in June in Piazza del
Duomo.
It would, however, be too simplistic to say that the
Cimento disappeared in 1668 with the emigration of
Leopold de’ Medici. In reality, the young people
continued to meet and to undertake new research.
These events were witnessed by Pietro Paolo da Sangallo, who had closely studied Redi’s Experiments
on the reproduction of flies, gnats, mosquitoes,
grasshoppers, butterflies, mites, scorpions, spiders
168
B. Baccetti - Malariology history in Italy
and finally, continuing the eternal argument with
Father Atanasio Kircher, on the generation of frogs
and vipers. Redi had discovered that all these creatures lay their eggs in suitable environments and
that from these emerged little vermiform animals
from which develop adults identical to those which
had produced the eggs. Aristotle and Pliny thought
that mosquitoes and flies were born in the slime of
certain worms that they called Ascaridi, and Ulisse
Aldrovandi (1602) also believed this. Ten years after
Redi studied mosquitoes in detail and bred them in
special well-sealed glass containers; da Sangallo
began similar experiments, on 20 June 1679, using
stagnant water from the plant nurseries located in
Florence at the beginning of the road which led to
Poggio Imperiale. He was inspired by the research
that Redi had conducted on flies and here he bred
mosquitoes. From these studies he recognised different species or varieties (like Redi, he used a
microscope); he drew good designs of all of the
important stages of metamorphosis and of the vari-
Fig. 2. In: da Sangallo PP, 1679, Tav I, drawing showing the
eggs (1), larva and pupa (2 and 3), and two adults (4 and
5) of the mosquito.
ous stages of life which demonstrated the necessity
of water in their life cycle.
This work was the subject of a report in the form
of a letter, in Tuscan dialect, dedicated to the most
illustrious Francesco Redi and printed in Florence
by Vincenzo Vangelisti, the Printer of the Archbishop, on 4 November 1679. The study is rich in illustrations which portray the different phases of the
biological cycle (da Sangallo, 1679, Table 1, Figs 1,
2, 3) and the two main forms of adults (Table 1,
Figs 4 and 5). One of these (Fig 5 id) looks very
much (not completely) like the adult mosquito
drawn ten years previously by Francesco Redi in his
Experiments (Redi, 1668, Table 29). On many other points, da Sangallo echoes the masterpiece of his
Master. He also cited and quoted the Arab naturalist Alchazuino (Zaccaria Ben Muhammed Ibn Mahmud) who had also described the mosquito, comparing it to an elephant with wings.
Da Sangallo ended his study with the words:
Fig 1. In: Redi F, Opere, Tomo I, Hertz, Venezia, 1742, Tav.
29, Zanzara. This table seems part of “Esperienze intorno
alla generazione degli insetti” where Mosquitoes are quoted in the text. In the Venetian edition this drawing is
arranged together with others ones showing “Pollini”,
“Pidocchi”, and other parasites.
“Per liberarsi da così fatta molestia delle Zanzare sono insegnati da diversi autori molti, e diversi, medicinali provvedimenti. Plinio loda l’ungersi ogni sera tutto quanto con l’olio d’assenzio. Lo stesso Emilio
Macro non sapendo per avventura che anche le Zanzare amano il vino, suggerisce di bagnarsi tutto con
questa bevanda, purché vi sia stato infuso e bollito
l’assenzio. Alcuni altri insegnano impiastrarsi la faccia, e le mani, e le braccia con la saliva dopo che s’è
ben bene masticato il cumino, e poscia si mescoli con
vin bianco potente, e fumoso e con esso se ne aspergano le finestre, e le porte, e tutta quanta la casa, e
questo lavoro per maggior complicazione si faccia con
ramoscelli fronzuti, e verdi. L’autore del libro de’
Medicamenti semplici a Paterniano attribuito a Galeno vuol che si adopri il sugo de’ frutti della Tamerigia, ovvero la loro decozione fatta in acqua. Altri
lodano il bagnarsi il capo e tutto quanto il corpo con
la bollitura di Ruta, o di Nigella, o di Coniza, aggiuntovi ancora per maggiore efficacia una buona quantità di vetriolo, e di carboni di ginepro, il che mi
immagino, che faccia un bel vedere. Vi è chi propone
impiastricciarsi ogni sera tutto quanto da capo a piedi nell’andare a dormire con un certo guazzabuglio
fatto d’olio, d’aceto e di salvia pesta, e se ad alcuno
non piacesse la salvia, vi è chi in suo cambio pone la
polvere dell’incenso. Quei Greci che scrissero sull’agricoltura approvano per cosa utile circondare il letto
con una ghirlanda fatta di fronde di canapa, che sia
stata spruzzata d’acqua, ed un certo valent’uomo propone che si tengano in vicinanza del capo e sotto le
piante dei piedi spugne inzuppate nell’aceto forte, e
che una simile spugna s’attacchi nell’alto della casa,
e quello che mi pare più considerabile, o per dir
meglio ridicolo, si è, che volendo la ragione, per la
quale sia giovevole così fatta spugna attaccata nell’alto della casa, dice che le Zanzare correranno tutte
a svolazzare intorno a quella spugna colassù appiccata, e non s’avvede che se ciò sarà vero elle voleranno ancora intorno al capo e intorno ai piedi di
colui che avrà messo in opra così prelibato consiglio.
Certuni ricorrendo alla simpatia, o all’antipatia delle
cose, o per dir meglio alla superstizione, scrivono che
lo attaccare nel bel mezzo della casa un pelo di Cavallo sia rimedio infallibile contro il ronzio, e contro le
punture delle Zanzare e forse credono costoro che sia
vero che Apollonio Tianeo co’ suoi incantesimi operasse (come racconta Tzeze) che nelle Città
d’Antiochia e di Costantinopoli non entrassero mai
vive le Zanzare. I suffumigi, che a questo fine vengon
proposti dagli Autori son tanti, e tanti, che io per me
credo, che tanti non ne sapessero, e non ne mettessero in esecuzione il Mago Ismeno, e le Fate del Boiardo, e dell’Ariosto. Tutte queste baie, ancorché tenute
per vere dal credulo volgo, sono totalmente inutili, e
fastidiose, e moleste più delle Zanzare istesse, contro
le quali un bel riparo mi sembra quello solo, ed unico, che fu ritrovato anticamente da’ pescatori dell’Egitto, cioè a dire un buono Zanzariere, che perfettamente circondi il letto, e a’ nostri tempi sia fatto di
gentilissimo velo di Bologna”.
To liberate oneself from the nuisance of the mosquito
has been the concern of many, and different medical
measures have been recommended by different
authors. Pliny suggested oiling oneself all over every
evening with wormwood oil. Emilio Macro had suggested the same, and not knowing that mosquitoes
also love wine he suggested bathing with this beverage, provided it had been infused, and the essence
boiled. Some others suggested smearing the face,
hands, and arms with saliva after chewing cumin, and
then mixing the saliva with strong white wine, and
sprinkinge windows, doors, and the whole house with
themixture – an activity which should be doen with
169
leafy green branches. The author of a book of simple
medicaments of Paterniano attributed to Galen,
advised using the juice of tamarisk fruits, or rather
their decoction in water. Others commend wetting the
head and the whole body with boiled ruta, or nigella,
or coniza, adding to them for still more efficacy a
good quantity of vitriol, and of carbonised juniper,
which I imagine would improve one’s appearance.
There are those who propose smearing oneself completely from head to foot every evening when going to
bed with a certain mess of oil, vinegar, ground sage,
and if one doesn’t like sage, there are those who suggest replacing it with incense powder. Those Greeks,
who wrote about agriculture, approved the usefulness
of surrounding the bed with a garland made from
hemp fronds, which was sprayed with water, and a
certain clever man proposes that you keep sponges
soaked in strong vinegar near your head and under
the arches of the feet, and a similar sponge fastened
high up in the house, saying that the mosquitoes will
all rush to fly around that sponge hung high – not
realising, if this is true, that they would also fly
around the head, and around the feet of whoever had
followed this delicious advice. Certain people writing
from sympathy or aversion to this method, or rather
from superstition, record that hanging the hair of a
Horse in the middle of the house is an infallible remedy against the buzzing, and stings of mosquitoes and
maybe they believe, if it is true, that Apollonio Tianeo
with his spells worked (as told by Tzeze) that in the
cities of Antioch and of Constantinople mosquitoes
never entered alive. The fumigations, that for this reason were proposed by the Authors, are many, and
many, I believe myself, had not been tried, and the
Magician Ismeno, the Fairies of Boiardo and of Ariosto did not perform them. All these potions, still
believed in by the credulous common people, are
totally useless, and annoying, indeed they are more
nuisance than the mosquitoes themselves, against
which a good shelter seems to me to be one and the
only protection, as was discovered ages ago by the
fishermen of Egypt, that is to say a good mosquito net
perfectly surrounding the bed, and in our times made
of a very soft voile from Bologna.
That night the students talked together, citing Redi. I
would like to think that also Giuseppe Del Papa
(1648-1735), another very young student of Redi,
and also influenced at a young age by the Cimento
Academy, was among this group. In fact, in his book
on grasshoppers (recently reprinted by Baccetti et al.,
2005), he also cited Alchazuino, who evidently had
many readers in Florence among the former followers
of the Cimento. In this circle, besides da Sangallo, we
have found many names of students in various obituary notices of Redi that appeared at the end of the
1690s, following his death in 1697. Among these, the
most well-known was, and still is, Giuseppe del Papa,
who died in the 1700s. But Pietro Paolo da Sangallo,
after having printed the most interesting and entertaining pages of post-Cimento entomological literature, which appears deeply permeated by the collaborative spirit of the Florentine Academy, and having
clarified many fundamental points of the biology of
the mosquito, disappeared from the scene. No one
170
mentioned him again, except for Abbot Salvino Salvini, an illustrious Arcadian who in his obituary of Redi
in 1699 noted the famous letter on the mosquitoes
that da Sangallo had printed in Florence in 1679 and
dedicated to Redi. Salvini finally in 1684 published a
further paper (Florence, Piero Matini: Osservazioni
intorno agli Animali Viventi che si trovano negli Animali Viventi).
Even earlier, on 14 October 1690, in a letter to
Giuseppe Lanzoni, an illustrious physician in Ferrara
and the author of two esteemed manuals, Zoologia
parva (Ferrara, 1689) and De Balsamatione (Ferrara,
1692), Redi wrote: “It was a miracle, that I should
have found one of those letters that Pietro Paolo da
Sangallo wrote to me about the Generation of Mosquitoes. Whomever wants to pay 100 ducats, I do not
believe that one could find another, because as your
Excellency could see, it has been a long time since it
was printed, and this Doctor died shortly after it was
printed. The virtuous genius of your Excellency, and
very well deserving of good philosophy was the reason that I was able to find it. I send you in this letter
what you commanded me to send” (Redi, 1690,
p. 214). Pietro Paolo da Sangallo must have died at
the beginning of the 1690s. He was only a boy, the
youngest of the last followers of the Cimento, as he
described himself, he was also a doctor, as Redi also
described him. And no one has ever written about his
life, except for his Master. In the letter to Lanzoni,
Redi said: “I send you the Trattatello delle Esperienze intorno alla generazione delle Zanzare, printed by
Sir Pietro Paolo da Sangallo. Here in Florence nothing else has been written and printed about those
Mosquitoes” (Redi, 1690, p. 214).
A final comment on the brilliant correspondence of
Redi. Diacinto Cestoni, a pharmacist in Livorno
who worked with Redi on the origins of scab, was
one of his main correspondents. In 1691 Francesco
Redi wrote to him with great enthusiasm: “I can
give you the news that for some weeks my health
has been much, much better; ‘che ella duri’ ‘may it
last’, said Gian Bracone when he fell from the tower and saw in the air that he was not hurt; but that
the damage would be, when he hit the ground”. And
then Redi concluded “The Most Serene Grand
Duke 2 and the Most Serene Grand Duchess Vittoria3 wanted to read my letter and they were very satisfied with it” (Redi, 1691, p. 224).
In 1717, the Italian physician Giovanni Maria Lancisi published a treatise on swamp fevers in which
he suggested that marsh fever was due to some
marsh poison transmitted by mosquitoes.
In 1930 the American professor L.O. Howard,
one of the most eminent entomologists and author
of A History of Applied Entomology (Smithsonian
2
3
Cosimo III.
Vittoria della Rovere, the mother of the Grand Duke.
Institution), described the Tuscan Professor of Zoology Eugenio Ficalbi (1858-1922) as “a competent
entomologist, who had written about Mosquitoes
before they were proved to be carriers of malaria,
published in 1899 and 1901 important papers upon
Italian Culicids”. Twenty years before (1911), Battista Grassi (1854-1925), who was at that time the
star of Italian biology, including malariology,
declared of Ficalbi “...his papers concerning Culicids
initiated the modern research on the systematics of
these diptera, and greatly facilitated the experimental studies of Grassi, in searching for the insect
transmitting malaria” (Grassi, 1911, p. 123). In
fact, Ficalbi (Ficalbi, 1899-1901) published nine
papers on Italian Culicidae in Florence and in Siena
(1889-1896), as well as a systematic review of the
family in Europe, and a list of 20 Italian species
belonging to the group. In 1901 he also published
an other important study, Sopra la malaria e le Zanzare malarifere nella Salina di Cervia e nel territorio di Comacchio. The name of Ficalbi must therefore be included in the history of medical entomology – his contribution ensures that Italians played a
significant role in the history of the subject.
However, the most important name associated with
the malaria story is that of Battista Grassi, a man of
broad training, educated in Germany and married to
a German woman, who wrote extensively on many
zoological (intestinal worms, biology and reproduction of murenoids and eels) and entomological topics.
In this latter field, Grassi studied Diptera (Phlebotomus, Culicids), Aphis embryology, Diplura and Thysanura, Embioptera and Termites – studies that were
awarded (1898) the Darwin Medal of the Royal Society for Applied Entomology and Parasitology. He was
also the first to furnish proof that Anopheles claviger 4
is a carrier of malaria (1898), and in November of the
same year, together with Giuseppe Bastianelli and
Amico Bignami, he performed the first experimental
transmission of human malaria in Anopheles Mosquitoes, and observed (1899) the complete life-cycle of
the different species of human Plasmodia. The Plasmodia had first been discovered in human blood by
Alfonse Laveran in 1880. In 1898, Ronald Ross elucidated the entire life-cycle of bird malaria. This discovery was confirmed the following year in humans
by Grassi and colleagues, but only Ross was awarded
by the Nobel Prize for malaria (1902). Camillo Golgi
(University of Pavia) obtained the Nobel Prize in 1906
for the study of the fine structure of nervous cells, but
he had also demonstrated in 1886 that different
species of Plasmodium are responsible for two types
4 As a matter of fact the Italians did all their experiments
with a species of maculipennis complex presumably An.
labranchiae as demonstrated by Coluzzi M. and Corbellini G.,
1998, Il centenario della malariologia (1898-1998), Parassitologia 40(4):361-375 on the basis of Grassi’s drawing which
provided an excellent representation of the egg which is diagnostic for this species.
of intermittent fever (Plasmodium malariae and Plasmodium vivax). Moreover, in 1889 Golgi found the
link between the rupture of infected erythrocytes and
the onset of the fever. In conclusion, the Italian medical literature on the subject was very extensive. A
number of medical men, including E. Marchiafava, A.
Celli, G. Bastianelli and A. Bignami had been assiduously studying malaria and publishing for many years
before Anopheles was discovered to be the vector,
especially in the interval between the finding of the
causative organism of the disease by Laveran in 1880
and the eventful year 1898 when the mosquito relationship was discovered by Ross in avian malaria and
Grassi published the paper Relations between malaria and certain insects. A curious detail reported by
Howard in 1930 is that when he called Grassi in
Rome, in 1923, he sent him printed documents which
he claimed conclusively proved that Ross deserved
credit only as the discoverer of the vector of sparrow
malaria. Howard and Grassi became friends that same
year. In Howard’s book of 1930 we read: “It was on
this trip that Grassi showed me the interesting mating
of Anopheles at nightfall about certain pigsties on the
estate. He had been the first person to observe this
mating, in spite of the efforts of many men in many
countries for many years. Grassi’s work, especially in
this region had been systematic, and he showed me a
mass of records that had accumulated and which
undoubtedly contained many facts of value. His special interest in this region continued until the time of
his death. Since 1924 when the International Health
Board has stationed a representative, Dr L.W. Hackett, in Rome. But the bitter controversy between Grassi and Ross was carried on vehemently until Grassi’s
death in 1925. Howard, in his book of 1930, reports:
“even in 1927, when I called on Ross in England, he
could not speak of Grassi without profanity. “Celli”,
he said, “was a gentleman, but Grassi was a damned
pirate”. (Howard 1930, p. 491).
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Howard LO (1930). A History of Applied Entomology. Smithsonian misc Collection, 84, Washington, 545 pp.
Lancisi G (1717). Noxiis paludum effluvis eorumque remediis.
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Laveran A (1880). Note sur un nouveau parasite trouvé dans le
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3 CAMBEFORT:Layout 1
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Knowledge of Diptera in France from the beginning
to the early twentieth century*
Y. Cambefort
Laboratoire Rehseis, UMR 7596, Université Paris 7, France.
Abstract. Although insects have been objects of observation in French-speaking countries since the
seventeenth century, and were illustrated by Réaumur and other scientists during the eighteenth, specialized dipterology only emerged in the first half of the nineteenth century. The two main divisions of the
Order Diptera were defined by French entomologists, namely Nemocera (currently Nematocera) by
Latreille in 1817, and Brachocera (currently Brachycera) by Macquart in 1834. Insects as a whole were
rarely studied until the late nineteenth century, when the discovery of their role in the transmission of
important diseases resulted in the creation of a new discipline: medical entomology. From this time on,
medically important groups (mosquitoes, tsetse flies, etc.) have been objects of intense concern and
study, especially at the Pasteur Institute and the Paris Faculté de médecine. But the most important
French dipterist* in the twentieth century has probably been the Muséum specialist Eugène Séguy.
Key words: entomology, France, Diptera, mosquito, fly.
Mains chasseresses des diptères
Dont bombinent les bleuisons
Aurorales, vers les nectaires,
Mains décanteuses de poisons…
Oh! quel rêve les a saisies?
Arthur Rimbaud
Interest in insects manifested itself early in history.
Biting and noxious species have always been familiar, and are mentioned in the Bible on various occasions. Virgil’s Culex is a curious testimony of concern respecting these insects, to the point of building a tomb to one of them (but the story might have
been ironical). However, the group has been poorly
known, even unrecognized, up to relatively recent
times. The very word Diptera, introduced by Aristotle, was rarely used before Linnaeus. Only after
the invention of microscope were scholars able
properly to observe and identify these insects. Eighteenth-century systematists attempted to separate
and name genera and species among them. However, both large scale classification of the order
Diptera, and the subtle distinction and characterization of its almost innumerable species, did not significantly progress before middle nineteenth centuCorrespondence: Yves Cambefort, Laboratoire Rehseis, UMR
7596, Université Paris 7, Centre Javelot, 2 place Jussieu, 75251
Paris Cedex 05, France, e-mail: yvecambe@club-internet.fr
This paper combines and revises the two communications presented by the author at the Workshop on History of Medical
Entomology: “From cabinets of curiosities to entomological reference collections” (London, Wellcome Trust Centre, April 2223, 2005); “The History of Diptera systematics in France in the
nineteenth and early twentieth centuries” (Rome, Accademia
Nazionale dei Lincei, October 11-12, 2007).
* The term “Diptera” denotes single winged insects with biting
mouth parts. The word “dipterist” for one who studies these
insects does not exist in the English language, but has been coopted from the French for the purposes of this volume.
ry. This paper presents a general sketch of French
Dipterology, from its origins in seventeenth century
Europe to its development in the nineteenth and
twentieth centuries. In the first half of this history,
Dipterans were just difficult and rather unattractive
insects, studied by a few devoted specialists. However, a revolution took place from the 1880’s, when
it was realized that Dipterans played a significant
role in the transmission of major diseases in temperate and especially in tropical countries. Dipterology ceased to be anecdotal, and became a significant
component of medical and veterinary parasitology.
From curious to scientists
In the sixteenth and seventeenth centuries, princes
and prominent citizens used to assemble cabinets of
curiosities, which were private museums of a sort,
where two groups of items were preserved: natural
objects, or Naturalia, and artificial objects, artefacts
or Artificialia. Soon (ca. 1550), however, such a
cabinets became fashionable also among ordinary
educated people. One book published in 1565
advised such readers wishing to establish inexpensive cabinets to select cheap objects, like “small animals” (Quiccheberg 1565, in Mairesse 2004, p. 97)
– which designation also included insects. Jacob
Hoefnagel’s 1592 album contains remarkably accurate engravings of insects (including some thirty-five
Dipterans) carved after his father Joris Hoefnagel’s
iconographic models that must have been designed
from actual insect specimens, probably kept in cabinets (on Joris Hoefnagel, “the first Belgian entomologist”, see Leclercq 1987). The first two known
treatises on entomology (Aldrovandi 1602, and
Moufet 1634) are both profusely illustrated with
less accurate but generally recognizable woodcuts,
also certainly carved after actual models from cabinets or collections. Aldrovandi’s third book is devot-
174
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Y. Cambefort - Knowledge of Diptera in France
ed to Anelytris Bipennibus and divided into five
chapters: I, De Musca (pp. 342-371); II, De Musca
Vinaceorum (p. 372); III, De Ephemero (pp. 372373); IV, De Oestro et Tabano (pp. 373-381); V, De
Culicibus (pp. 382-402). The sixty pages contain
seventy-four woodcuts of flies and twenty-two of
mosquitoes, gnats, and midges, actual size, and generally of a good quality: the best that could be
achieved without a microscope. The text, however,
is disappointingly prolix and futile. It is worse in
Moufet (1634, 1658), where two- and four-winged
insects, from crane-flies to dragonflies, are intermixed in Chapter XI “Of the divers kindes of Flies”.
Chapter XIII, “Of Gnats”, which is more limited in
scope, and is not illustrated.
By the seventeenth century, cabinets of natural history retained only the naturalia component of the
former, larger cabinets of curiosities, and were widespread among middle-class citizens. At that time,
paintings of the so-called “still-life” and “vanity” genres often depicted flies, which always had a connotation of death (Chastel 1984; Doby 1999). I have
explained elsewhere how medicine and painting
jointly gave birth to scientific entomology (Cambefort 2004). Insect collections – which might be considered a reduced type of natural history cabinet –
are documented in France in the seventeenth century. For example, the famous British traveller John
Evelyn mentions the collection of the Parisian gardener and collector, Pierre Morin, called “le Jeune”:
“The next morning (April 3rd, 1644), I was had by a
friend to Monsieur Morines Garden; a person who
from an ordinary Gardner, is arrived to be one of the
most skillfull & Curious Person of France for his rare
collection of Shells, Flowers & Insects (...) The very
greatest curiosity which I esteemed, for being very
ingenious and particular, was his collection of all the
Sorts of Insects, especially of Buter flys, of which he
had so greate Variety; that the like I had never seene:
These he spreads, & so medicates, that no corruption
invading them he keepes in drawers, so plac’d that
they present you with a most surprizing & delightfull
tapissry” (de Beer 1955, t. 2, pp. 132-133).
These kinds of collections were, in fact, still made
for purpose of decoration and contained almost only
large and bright insects, especially butterflies. But
some collectors changed into authentic amateurs,
even scientists. One of the first and best examples is
Jan Swammerdam (1637-1680), Dutch physician
and one of the first authentic entomologists, who
produced works of a much higher quality than those
of Aldrovandi and Moufet. His first important
book on insects (1685) has been translated into
French, and it was probably eagerly sought after and
read by the first French amateurs. It contains
engravings of the larva, nymphs, and adults of Culex
(pp. 100-110, plates II and III). These engravings
are much better than those in Robert Hooke’s
Micrographia of 1665, which is quoted and discussed. In addition, Swammerdam gave a description of mosquito biology, which, together with the
engravings, became a standard – or rather a topos –
up to the nineteenth century. He also carefully distinguished male and female adults, especially by the
antennae and mouthparts. Later in the book various
other Dipterans (horse flies, drone flies, etc.). are
described and illustrated.
Eighteenth century entomologists
Natural history cabinets were fashionable among
French nobles and citizens alike. Abbé Pluche’s
Spectacle de la Nature (1732), one of the most
famous and influential natural history books of the
early eighteenth century, gives a picture of countryside aristocrats amusing themselves with observations on animals and plants (since nobles were not
supposed to work). The Count in the story possesses a cabinet “which assembles all the conceivable
species of animals” (p. 2). Flies and gnats are the
objects of the huitième entretien (eighth conversation), which takes place between Count, Countess,
Prior, and Knight. They quote, among others,
Swammerdam’s observations on mosquitoes (pp.
204-209). Detailed engravings of mosquito mouthparts are also taken from Swammerdam.
Réaumur
True scientists also assembled natural history cabinets, and one famous French gentleman must be
mentioned here: René Antoine Ferchault, seigneur de
Réaumur (1683-1757). He is well known for a number of inventions in physics and technology, but his
most important scientific works are probably the
Mémoires pour servir à l’Histoire des insectes, six
volumes of which were published between 1734 and
1742. In the fourth volume (1738), third Mémoire,
Réaumur gave a summary of Diptera classification:
De la distribution générale des Mouches, en classes,
en genres & en espèces (pp. 123-161 and plates 811). But it is rather deceptive, since it starts by making a division between two-winged and four-winged
“flies”! Clearly, the author was not interested in systematics, and was able to recognize only the most
obvious groups. But this superficial interest in systematics is more than compensated for by the exquisite acuteness of the biological observation. In the
fourth volume, nine Mémoires are devoted to various
species of flies, especially house flies, so-called bluebottles and greenbottles, and other carrion feeding
flies. The twelfth Mémoire contains important observations on sheep nostril-flies (Oestrus ovis) which
are among the first published on this species. Then
came the thirteenth and last Mémoire of the volume,
the “Histoire des cousins” (cousin was the name given at that time to modern “moustiques”, in English
“mosquitoes”). In this classic of entomology, Réaumur described in a lively, almost empathic style, the
behaviour and metamorphoses of the mosquitoes he
was able to observe in the French countryside. Réaumur’s text is more detailed and as a whole much
more interesting than Sammerdam’s. He had also
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some nice engravings made, perhaps not better than
Swammerdam’; and was good enough to be referred
to by Linnaeus: the figure 3 on Réaumur’s plate 43
of the fourth volume was cited in the tenth edition
of Systema Naturae (1758), in the paragraph devoted to Culex pipiens, and for this reason has been
selected as the “lectotype” of this species (Harbach
et al. 1985). From Réaumur’s plate 40, in the same
volume, the engraving of the adult male (figure 2)
was selected by Harbach et al. (op. cit.) as the “lectotype” of Culex bifurcatus Linné, 1758 (a species
often confused with Anopheles bifurcatus Meigen,
1818). Réaumur’s fifth volume (1740) starts with a
Mémoire devoted to crane-flies, a sort of hold-all
group which contains every gnat and midge which
are not mosquitoes. The second Mémoire gives interesting accounts of deer nostril-flies (Oestridae),
which supplement those on sheep nostril-flies in the
fourth volume.
Geoffroy and Fourcroy
Réaumur’s sixth and last volume was published in
1742. Twenty years later, an anonymous, two volume
book was published in Paris: Histoire abrégée des
Insectes qui se trouvent aux environs de Paris. Its
175
author was a Parisian physician, Étienne-Louis Geoffroy (1727-1810), and the book was reissued in 1764
under his name. It is one of the first treatises on
insect systematics. However, as it does not use the
binominal Linnaean system, it has posed a number of
taxonomic problems. In general, all its new genera
are considered valid (e.g. the stable-fly, Stomoxys),
but its species are not. Diptera are dealt with in the
second volume (pp. 430-580). The author clearly
indicates the key character of the group: the presence
of two wings, and names these insects in French and
Latin: Diptères and Diptera. A table is given of the
thirteen genera which are recognized and could be
identified from morphological characters, especially
antennae and mouthparts (Fig. 1).
The table is not arranged in a dichotomous order:
the two main divisions within the Order Diptera
have not yet been individualized, and the Order is
rather perceived as forming a “ladder” (scala naturae), according to the conception of the time (see
e.g. Daudin 1927). In all, 175 species are described.
Geoffroy’s collection is still in existence in the Paris
Muséum (Cambefort 2006); but the Diptera have
almost disappeared. Some years after Geoffroy’s
book, a summary with a few complements was pub-
Figure 1. Étienne-Louis Geoffroy’s Histoire abrégée des insectes des environs de Paris (1762), table of Diptera genera.
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Pagina 176
lished by the chemist and biologist Antoine François
de Fourcroy (1785). This time, the Linnaean binominal system was followed throughout.
[see a chronological table of chairmen of entomologie at the Paris Muséum in Table IV].
The Société entomologique de France
Baumhauer and Meigen
In the last years of eighteenth century, a German
amateur entomologist spent some his time in Paris:
Johann Matthias Baumhauer (1759-1818). He had
assembled a large collection of Diptera (parts of
which are still preserved in the Liège and Leiden
museums), and vainly tried to study and identify
them, since at that time they were poorly known.
But one of his friends (some say a relative) had
simultaneously begun to study the group: Johann
Wilhelm Meigen (1764-1845), later called “the
father of Dipterology” since he was the first to publish an extensive monograph on these insects (18181838). But before 1800, Meigen was a mere beginner, whom Baumhauer wished to encourage. He
sent part of his collection to Meigen, who worked
on it for many months, and finally sent it back to
Baumhauer together with detailed comments.
Baumhauer edited this text, gave it to the Parisian
bookseller Fuchs, and it was published as a booklet
of some forty pages (Meigen 1800). It was Meigen’s
first published work, giving a synopsis of his whole
future system. Unfortunately, he later changed certain names, which complicated Diptera systematics
until the 1800 booklet’s priority was cancelled by
the International Commission of Zoological Nomenclature.
Nineteenth century specialists
Latreille
Réaumur, Geoffroy and Fourcroy were physicians or
surgeons or engineers, etc. Baumhauer and even
Meigen were mere amateurs. But Pierre-André
Latreille (1762-1833) was the first true professional
engaged in the study of Entomology (Dupuis 1974).
In 1798, he joined the Muséum d’Histoire naturelle,
in Paris, to sort and arrange the insect collection, but
it was not until 1830, only three years before his
death, that a new chair was created for him at the
Muséum: “Zoologie des Crustacés, Arachnides et
Insectes”, or “Animaux articulés”. In 1802, in the
famous series of Sonnini’s Suites à Buffon, Latreille
established for the first time, under the name Tipulariae, a group of genera with long, multiarticulate
antennae (Tipula and Culex of Linnaeus), and separated them from other Diptera. Later (1817), he gave
this group a family rank and called it Némocères
or Nemocera. It was the first conception and
name of the current Nematocera, or “long-horned
flies”. In 1825, Latreille divided his Némocères into
tribes and created the tribe Culiciformes: this was the
first time mosquitoes were given a family rank;
Latreille’s chair, at the Muséum, was continued until
1997, with 10 professors succeeding him, but only
one of them being a dipterist: Eugène Séguy (below)
In February 1832, the Société entomologique de
France (hereinafter “S.E.F.”) was created, the oldest
of national entomological societies, and Latreille
was elected its honorary president (he would die the
following year). At the end of 1832, there were in
all 94 members in the S.E.F., only four of whom
were professionals: Latreille, his aides-naturalistes
Jean-Victor Audouin (who succeeded Latreille in
1833) and Auguste Brullé, and his préparateur Hippolyte Lucas. A few more members, who belonged
to the Muséum staff and were elected because of
their fame (e.g. Cuvier, Geoffroy Saint-Hilaire),
might be considered as professionals, but hardly as
entomologists. Henri Milne-Edwards was not yet an
entomologist, but he would become one when he
succeeded to Audouin in 1841. In all, about ninety
percent of the S.E.F. members were pure amateurs,
and mostly collectors: as in the eighteenth century,
they were interested almost exclusively in the largest
and most brilliant insects (butterflies and beetles),
and they rarely showed any scientific interest in the
objects of their passion. However, a few of these
amateurs changed into true scientists; they worked
and published on anatomy, biology, etc.: the name of
Léon Dufour (1780-1865), author of important
monographs on insect anatomy, must be mentioned
here (e.g. Dufour 1851). As for dipterists, there was
only one among the S.E.F. members of 1832: Justin
Macquart, and only four more were admitted in the
following thirty-six years: Jean-Baptiste RobineauDesvoidy (in 1833), Jacques Bigot (in 1844), Louis
Pandellé (in 1850), and Émile Gobert (in 1868).
After them, the next members will be dealt with in
the following section.
One observation should be made here: there have
always been close relationships between the
Muséum’s entomology department and the S.E.F.
For example, although professional entomologists
have always been a minority compared with amateurs, the S.E.F. president was alternately a professional and an amateur. Further, members of the
S.E.F. have always had free access to the collections
of the entomology department.
Macquart
Justin Macquart (1778-1855), from Northern
France, was the first French specialist in Diptera.
During all his life, he travelled extensively across
Europe, especially in the German-speaking countries
(he gives a narrative of his life in Macquart 1850).
As he was fluent in German, he could exchange an
important correspondence with Meigen and Wiedemann (see below). His early works was directly
inspired by Meigen’s first volumes (1818-1838),
four of which were published at that time. In his
first monograph (Macquart 1826), he clarified the
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177
Table I. Dichotomous table of Musciae [sic] (flies s. str.) in Macquart (1843), p. 268 bis.
distinction made by Latreille between Tipulaires or
Némocères (or Nemocera) and other Diptera. For
Macquart, all Diptera should be divided into two
groups: Némocères on one hand, and the rest on the
other. He did not give the latter group a name until
a few years later, when he coined the name Brachocères, symmetrical to Latreille’s Némocères
(Macquart 1834, p. 183). It was the first time the
classic, dichotomous division of the order Diptera
into two suborders had been formalized.
Macquart’s first works on Dipterans attracted the
attention of the Paris Muséum administrators, and he
was asked to study the institution’s collection. In
1839, he went to Meigen’s home in Stolberg (near
Aachen), and managed to have Meigen’s collections
and documents acquired by the Muséum (Macquart
1847), including a magnificent series of 305 original
watercolours, with more than 4000 individual illustrations depicting the habitus and details of all the
European Diptera identified by Meigen during almost
fifty years of continuous activity (these plates were
published only thirty years ago: Morge 1975-1976).
Meigen’s collections and documents helped Macquart
with his work in progress. But Meigen had been
working only on European Diptera, and Macquart
wished to work also on “exotic” species, especially
because the Muséum possessed and received large
numbers of the latter. For this part of his work, Macquart took his inspiration from another German
scholar, Christian Rudolf Wilhelm Wiedemann (17701840), who had published the first book on nonEuropean Diptera (Wiedemann 1828-1830). In his
1834-1835 monograph, Macquart tried to give as
complete a study as possible of French Diptera, men-
tioning only the types of exotic genera that were
known to him. Later, from 1838 on, he endeavoured
to supplement Wiedemann’s work on extra-European
Diptera. In the last part of his life, he worked almost
full time on the extensive material the Muséum
received from all over the world, describing 140 genera and some 2000 species of non-European Diptera
(Macquart 1838-1855).
Robineau-Desvoidy
Jean-Baptiste Robineau-Desvoidy (1799-1857) was
a physician, and still a young man, when he published his first work on mosquitoes, a short paper
(1827), but also when he published his much more
ambitious monograph on flies of the world, with its
more than 800 pages (1830). Only three years later
was he admitted to membership of the S.E.F.
(1833). He continued to study Diptera up to his
death, and his last monograph was posthumous
(1863). His activity as a dipterist is somehow symmetrical with Macquart’s: with the exception of his
first short paper on mosquitoes, he began his work
with an extensive revision of mostly exotic species,
using all the collections available in France, including the Muséum’s. Afterwards, when Macquart was
himself working on the Muséum’s materials,
Robineau-Desvoidy concentrated on European
species, and his last work returned to the environs
de Paris, the same words used by Geoffroy a century before; but Geoffroy’s 150 pages on Diptera were
then expanded into more than 2000 pages.
For his monograph on flies, Robineau-Desvoidy
was able to study the collection of one of the most
famous French amateurs: General Count Auguste
178
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Figure 2. Plate 14 from Justin Macquart’s Diptères exotiques nouveaux ou peu connus, Tome II, 3e partie (1843).
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Dejean (1780-1845). Dejean was more interested in
beetles, and he published an authoritative catalogue
of the Coleoptera order. But he had a large collection
of other insects as well, including Diptera, which he
had assembled during twenty years (1815-1835) by
buying up everything of any interest which came
onto the entomological market. He thus acquired a
collection brought in 1816 from the Congo by the
British traveller John Cranch. In this material,
Robineau-Desvoidy found a curious fly which he
mentioned briefly in his 1827 paper under the name
Némorhine (p. 396). In his 1830 monograph, he
more formally created a new genus and new species
for this fly: Nemorhina palpalis; he thought that the
fly’s proboscis was too weak to bite, and called it
“innocent” (Robineau-Desvoidy 1830, pp. 389-390).
Even if this 1827 mention is not taken into account,
the formal genus and species names were introduced
in a publication dated 30 June 1830. It was only on
22 September 1830 that Wiedemann’s second volume was published, which contained the description
of a new genus and new species: Glossina longipalpis (pp. 253-254). Clearly, Wiedemann’s genus
was the same as Robineau-Desvoidy’s (and possibly
also the species was identical). Five years later, in his
general monograph, Macquart considered RobineauDesvoidy’s Nemorhina as a synonym for Wiedemann’s Glossina, and the two species as identical
(Macquart 1835, pp. 244-245, pl. 16, fig. 8); he did
so also in his work on exotic species (Macquart
1843, pp. 269-271, pl. 14, fig. 1) (Fig. 2). In this latter work, he explained – like Robineau-Desvoidy –
that Glossina’s proboscis was too weak to pierce a
mammal’s skin and that the species feeds on flowers:
at that time, the first accounts of African travellers
had not yet been written, and the very name “tsetse”
was still unknown.
It is, however, difficult to explain why Macquart
took such taxonomic decisions, which would formally and firmly establish Wiedemann’s name for
the genus: was he unaware of the (however short)
priority of Nemorhina? Or was it because he did not
appreciate Robineau-Desvoidy? Austen (1903), in
the role of “first reviser”, did not rectify this mistake
(his explanations are rather embarrassed: see e.g.
his p. 51)1, and the genus-name Glossina was from
that time definitely established. But Austen recognized the validity of Robineau-Desvoidy’s species.
Today, Nemorhina Robineau-Desvoidy is generally
considered as a subgenus of Glossina Wiedemann,
with four species: caliginea Austen 1911, fuscipes
Newstead 1910, pallicera Bigot 1891, and tachinoides Westwood 1850, in addition to palpalis
Robineau-Desvoidy. This last species is one of the
most important vectors of sleeping sickness in Western and central Africa, and was therefore the sub1 In his 1903 book, Austen made another remarkable mistake: his title page attributed the name Glossina to the English entomologist Westwood, instead of the true author, the
German Wiedemann.
179
ject of particular study by French scholars in the
early twentieth century (see below).
Bigot, Pandellé, Gobert
The last dipterists to join the S.E.F. before 1870 were
Jacques Bigot (1808-1893), Louis Pandellé (18241905), and Émile Gobert (1838-1922). Bigot published a number of works on Diptera, especially on
extra-european species. In addition to a short revision of the genus Glossina (Bigot 1885), he later
described another West African species of this genus:
G. (Nemorhina) pallicera Bigot 1891. His collection
was sold to George Henry Verrall (1855-1911), and
is now preserved in the Oxford University Museum
of Natural History. Pandellé published important
monographs on French house-flies and their kindred
(Muscidae s. lat.); Gobert, originally a coleopterist,
became interested in Diptera later in his life, and
published some papers on these insects, including a
catalogue of French species (1887). Pandellé’s and
Gobert’s collections were donated to the S.E.F.,
which deposited them at the Paris Muséum in 1930.
Jean-Henri Fabre
Fabre (1823-1915, member of the S.E.F. in 1858),
was not a dipterist; but he must be mentioned here
since he has been one of the most influential entomologists ever, both in France and abroad. His
international fame was established by his Souvenirs
entomologiques, ten volumes of which were published late in his life (Fabre 1879-1907, Cambefort
1999). Here he intermixed observations and experiments on insect life with personal memories. Insects
were not presented in systematic order, but rather in
the order they appeared under his eyes, and he
always commented on their behaviour in a rather
subjective, even anthropocentric way. This was not
really science, but the style made the volumes lively, sometimes poignant, and – more important as far
as entomology was concerned – contributed to
attracting insects to the attention of a very broad
and wide public, almost all over the world. In the
first half of the twentieth century, Fabre’s fame
helped to raise funds for entomology (e.g. Bouvier,
in Ranc 1926). Fabre’s English translation, by
Alexander Teixeira de Mattos, did not follow the
original version’s lack of structure, but divided the
Souvenirs into fourteen taxonomically defined volumes, including one on Diptera (Fabre 1913). As
the order was Fabre’s least favourite, the volume
might have been shorter than average: it was completed with most personal memories, thus making it
– possibly contrary to Fabre’s intention – one of the
most charming of the series.
The emergence of Medical Entomology after 1880
and the three French circles
The demonstration by Patrick Manson in 1887 that
mosquitoes were involved in the transmission of
particular diseases marked the beginning of medical
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180
entomology. From this date on, dipterology changed
its status, from a rather abstruse and fanciful discipline to one of the major components of epidemiology. Everywhere in the world, but especially in
France, dipterists have always been few, as is
demonstrated by their small number in the S.E.F.
(see above) and their absence from the Muséum’s
staff up to 1918 (below). But their number then
began to increase in the 1890’s (Table II)2.
Table II. Respective numbers of dipterists received at
the Société entomologique de France between 1832
and 1931 (in successive periods of twenty years).
Period
Number of dipterists
1832-1851
4
1852-1871
2
1872-1891
2
1892-1911
16
1912-1931
17
If we consider the specializations of the S.E.F.
members for the period 1872-1931, we see that
fourteen of thirty-five dipterologists (40%) were
interested in medical and veterinarian dipterology
(Table II). But there is a bias in these small statistics: certain dipterists have never been members of
the S.E.F., and this is the case for some of the most
important actors of medical dipterology, e.g.
Alphonse Laveran, Félix Mesnil, Maurice NeveuLemaire, and Émile Brumpt. In fact, medical
dipterology from that time on was larger than the
other specialities. For this reason, the present section will not be divided into the specializations in
Table III, but will focus on medical dipterology.
Table III. Numbers of dipterists received at the Société
entomologique de France between 1872 and 1931
according to their specializations (n = 35).
Specialization
Number
General dipterology
16
Medical and veterinarian dipterology
14
Agricultural dipterology
5
The first dipterist who ever suggested that flies
might be involved in the transmission of diseases
was a military veterinarian, Pierre Mégnin (18281906), member of the S.E.F. in 1875 (Mégnin
1875). He later worked on the use of Diptera in
forensic medicine (Mégnin 1894), a field which he
created and which has been active up to now. But
apart from a few isolated personalities like Mégnin
and Fabre, most French dipterists were linked to
2
In fact, the total number of members of the S.E.F. also
increased, but much more slowly. For example, numbers
approximately doubled between 1852-1871 and 1892-1911,
while the number of dipterists increased eightfold.
one of three “circles” (or “schools”): the Institut
Pasteur, the Faculté de médecine, and the Muséum
d’histoire naturelle, which interacted with each other to a greater or lesser extent.
The Institut Pasteur
In 1880, the military doctor Alphonse Laveran
(1845-1922), working in Algeria, discovered the
parasite of malaria (genus Plasmodium). In 1884,
he hypothesised that mosquitoes were involved in
the transmission of malaria. In 1897, he joined the
Pasteur Institute, where he associated with the biologist Félix Mesnil (1868-1938). Both men worked
on blood parasites: first Plasmodium, and later Trypanosom. The latter had been known since the
1840’s, but was recognized as a major parasite of
man in 1901 (Laveran & Mesnil 1904, 1912). Neither men were entomologists (although Laveran
described a few species of exotic mosquitoes); but
they tried, and recruited assistants to help them in
their study of insects. Two of these must be mentioned here: Sergent and Roubaud.
Edmond Sergent (1876-1969, member of the
S.E.F. in 1905) had been working with Laveran and
Mesnil since 1900. He also attended Bouvier’s
course of entomology at the Muséum (below). In
1909, he published (probably with the help of his
brother Edmond) a practical handbook on bloodsucking insects (Figs. 3 and 4).
Sergent was appointed director of the Algiers
Institut Pasteur in 1910, and he spent some forty
years there studying various aspects of parasitology
and making significant discoveries (Sergent 1964).
Some of his students were also distinguished dipterists and produced important monographs (e.g. Sénevet 1935).
Émile Roubaud (1882-1962) also studied entomology with Eugène-Louis Bouvier at the Muséum, and
always maintained good relations with the Muséum
and S.E.F. (member in 1906, he was president of the
S.E.F. in 1927). Especially interested in parasitology,
he began his personal work with research on blackflies (Simuliidae). When Laveran and Mesnil began
work on sleeping sickness, they looked for a young
and talented entomologist, and Bouvier recommended Roubaud, who was almost immediatly recruited.
The famous Mission d’étude de la maladie du sommeil took place at that time (Martin et al. 1909):
from 1906 to 1909, Roubaud studied the biology and
ecology of tsetse flies in the French Congo, writing
his doctoral dissertation on the principal vector of
the disease in that country, which turned out to be
Robineau-Desvoidy’s “Némorhine”: Glossina
(Nemorhina) palpalis (Roubaud 1909). Later,
Roubaud associated with Bouvier in the anecdotal
“Zaharoff foundation for the study of flies” (see
below). More importantly, he founded (1914) and
for forty-four years directed, a laboratory for Medical Entomology and parasitology at the Institut
Pasteur of Paris, which served as a model for the
other Pasteur Institutes all over the world. In the
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181
Figures 3-4. Diagrammatic figures explaining Diptera taxonomic characters, in Sergent (1909), pp.
41-42.
3
4
same time, he began giving lectures on medical entomology at the Institute, but did not pretend to
replace the general entomology lectures which where
given at the Muséum. These structural innovations
were of the utmost importance: it can be said that,
at least up to the second half of the twentieth century, every French medical entomologist active in
France or in the French colonies, was trained by the
Pasteur Institute (as well as by the Muséum, as far
as general entomology was concerned). This resulted in the publication of a long series of works on
medical entomology, some of the shorter appearing
in the house journals: Annales and Bulletin de l’Institut Pasteur, and Bulletin de la Société de patholo-
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gie exotique. In addition to its permanent staff, the
Institute sometimes made use of collaborators. For
example, Father Jean-Jacques Kieffer (see below)
was sent in Algeria in 1922-1923 (probably at Sergent’s request) to study parasite Diptera.
monograph on sandflies (Larrousse 1921). In 1910,
Brumpt himself wrote a famous Précis de parasitologie which was repeatedly reissued until 1949
(Brumpt 1910).
The Muséum d’histoire naturelle
The Faculté de Médecine
In 1889, the professor and chairman of Histoire
naturelle médicale at the Paris Faculté de médecine,
Raphaël Blanchard, was admitted to the S.E.F. In
that year he published the first volume of an important treatise on medical zoology. In the second volume (1890), the Diptera section was deceptively
short. Mosquitoes were rushed through in four
pages, and Ross and Laveran were not mentioned.
However, in the fly chapter, Mégnin’s suggestion
was recalled: “Tsetse fly is harmless by itself, and it
is to be feared only because it propagates and inoculates the germ of a virulent sickness” (Blanchard
1890, p. 508, my translation). It is likely that Blanchard soon regretted this concision. Ten years later,
he was a member of the French Commission for the
study of malaria (Blanchard 1900), and published a
paper on mosquitoes of the Paris area (Blanchard
1901). Lastly, four years later again, he published a
large and fine volume on mosquitoes (Blanchard
1905). This book was much used among French
speakers, although it contains almost nothing original in comparison with Frederic Vincent Theobald’s
monograph on mosquitoes, then in progress, and
was completely superseded as soon as publication of
the latter monograph was completed in 1910. In the
meantime, Blanchard created the journal Archives
de parasitologie humaine et comparée in 1898,
where many papers on parasite insects were to be
published. In 1906, the title of his chair was
changed to Parasitologie. In these years, Blanchard
was much involved in zoological nomenclature; he
was also president of the Société zoologique de
France. He died in 1919, aged 62 and not yet
retired, and was replaced by Émile Brumpt (18771951), a great professor and a great parasitologist,
but not really an entomologist (he was not member
of the S.E.F.). Although Brumpt worked more closely with the Pasteur Institute than Blanchard, he created with two collaborators (Maurice NeveuLemaire and Maurice Langeron) a second journal in
1923 to counterbalance the Pasteur’s journals: the
Annales de parasitologie humaine et comparée.
Among his collaborators, Neveu-Lemaire – although
not a member of the S.E.F. – did interesting work on
sandflies (in 1906, he described Phlebotomus
dubosqi, now recognized as a significant vector of
leishmaniasis in Africa), and several mosquitoes
(Neveu-Lemaire 1923); later, he produced a volume
on medical entomology (Neveu-Lemaire 1938).
Langeron too was not an entomologist, but made
entomological collections in Tunisia (a Culicoides
langeroni was described by Kieffer in 1921). A third
collaborator, Fernand Larrousse, published a short
During the entire nineteenth century and up to
1918, there was no dipterist at the Muséum. In the
1900’s, when Austen was preparing his tsetse book,
his correspondent in the Paris Muséum was Joanny
Martin, who was in charge of Lepidoptera and some
other additional groups, including Diptera. However, as soon as he was appointed museum professor
of entomology in 1895, Eugène-Louis Bouvier
(1856-1944) expressed his interest in parasitology.
In the 1900’s, his lectures were attended by “pastorians”, and were even formally recognized from
1906 as part of the Pasteur Institute’s teaching, until
Roubaud began his own course in 1910. Bouvier
was been one of the organizers of the Mission
d’étude de la maladie du sommeil (see above), and
wrote, with Giard, the Instructions zoologiques (in
Martin et al. 1909; see also Bouvier et al. 1906).
Later, Bouvier always maintained – up to his retirement in 1931 – the best possible relations with the
Pasteur Institute, being member of various of its
committees. However, it may be asked why Bouvier
waited more than twenty years to recruit a dipterist
to the Muséum. Here are two partial answers.
First, the entomology section in the Muséum
always cooperated with members of the S.E.F. (see
above). In the years 1890-1914, there were some
good amateurs of Diptera: Father Jean-Jacques Kieffer (1856-1925, S.E.F. 1893), already mentioned,
who worked on midges and gnats, occasionally on
parasites (Culicoides), and wrote at the end of his life
the Faune de France volume on these groups (Kieffer
1925); Henry Brölemann (1860-1933, S.E.F. 1894),
who began work on mosquitoes, before changing to
millipedes; Dr Joseph Villeneuve de Janti (18681944, S.E.F. 1896), author of a long series of papers
on flies, especially parasites of other insects
(Tachinidae), and whom Bouvier ackowledged in
1923 as the leader of “almost all” French dipterists;
a later amateur can also be mentioned: Jacques
Hervé-Bazin (1885-1942, S.E.F. 1909), specialist of
hover-flies (Syrphidae), and better known for the
portrait his son Hervé Bazin drew of him in his novel Vipère au poing (“Viper in the fist”, 1948); his
important collection was offered to the Muséum.
Second, there was from 1895 in fact a dipterist,
not exactly “at” the Muséum, but “close to” it (près
de): Dr baron Jacques Surcouf (1873-1934), chef
des travaux de zoologie at the Laboratoire colonial
of the École pratique des Hautes-Études, “close to”
the Muséum. Surcouf, a member of the S.E.F. since
1905, worked particularly on horse flies (Surcouf
1909, 1924). He was also interested in South American Diptera, and one of his best works was a joint
venture with a Venezuelan colleague (Surcouf &
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Pagina 183
Gonzáles-Rincones 1911). But he was a difficult
man (perhaps due to his corsair descent!), and Bouvier could not rely upon him (Bouvier’s reservation
“almost all French dipterists”, above, was probably
an allusion to Surcouf).
Be that as it may, the recruitment of a dipterist
“at” the Muséum was necessary, and Bouvier looked
for a good candidate. He eventually managed to
find one: Eugène Séguy (1890-1985). Born to a
family of artists and trained as a painter, Séguy
began to become interested in Diptera in 1908,
more especially thanks to Villeneuve de Janti (and
to Surcouf, who however did not like Séguy’s
arrival in “his territory”). Because of the war, Séguy
was not recruited by the Muséum before 1918, as a
mere préparateur; but after that he became successively assistant, sous-directeur, and finally chairman
in 1956-1960 (Dupuis & Matile 1985; Haenni in
Séguy 2004). He was to live twenty-five years more,
only gradually decreasing his activity. Séguy published a long series of important works, the first
devoted to mosquitoes and other parasite Diptera,
as if Bouvier wished to make known that a void in
the Muséum’s expertise was now filled (Séguy
1923a, 1924) (Fig. 5). Séguy later published various
volumes in the series “Faune de France”; he was less
interested in the extra-European fauna, considering
that European species were still poorly known
(Séguy 1923b, 1925, 1926, etc.).
Incidentally, the Fondation Zaharoff pour l’étude
des mouches (“Zaharoff Foundation for the Study
of Flies”) should be mentioned here. In his continuous search for funding, Bouvier got in touch in the
years 1921-1923 with the famous arms dealer Sir
Basil Zaharoff (1849-1936), and persuaded him to
create this foundation. It did not last long, however,
and only few contributions were published from it:
short papers by Roubaud on the common house-
183
Figure 5. Cover of Eugène Séguy’s first book (Paris, 1923).
fly; by Pierre Lesne – who was a specialist in beetles, not of flies (Cambefort 2006) – on the lesser
house-fly; and by Surcouf on stable-flies. A more
ambitious contribution was announced (Villeneuve
de Janti on Tachinidae); but it was published later
and in another context. The foundation’s most
important production was Séguy’s volume in the
Table IV. Chairmen of entomologie at the Paris Muséum.
Jean-Baptiste de Monet de Lamarck (1744-1829) [chaire de zoologie des Insectes, Vers et animaux
microscopiques]
1793-1829
Pierre-André Latreille (1762-1833) [chaire de zoologie des Crustacés et des Insectes (crée pour lui
en 1830)]
1830-1833
Jean-Victor Audouin (1797-1841)
1833-1841
Henri Milne-Edwards (1800-1885)
1841-1861
Émile Blanchard (1819-1900)
1862-1894
Eugène-Louis Bouvier (1856-1944) [1917: division de la chaire en deux: entomologie s. str. et zoologie des Arachnides et Crustacés]
1896-1931
René Jeannel (1879-1965)
1931-1951
Lucien Chopard (1885-1971)
1951-1955
Eugène Séguy (1890-1985)
1956-1961
Alfred Serge Balachowsky (1901-1983)
1961-1974
Jacques Carayon (1916-1997)
1975-1985
Claude Caussanel (1932-1999)
1986-1997
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184
series Faune de France (Séguy 1923b), for which
Bouvier wrote a preface in which he discussed the
ins and outs of the foundation, and expressed his
sincere thanks to the generous patron (“généreux
mécène”) who financed it.
In the second half of his career, Séguy wrote more
general books, including a fine volume on Diptera
biology (Séguy 1950). He also found time to write
and publish elementary books, in order to attract
young people’s interest in the Diptera. Lastly, he edited a journal devoted to his favourite insects: Diptera
(11 volumes, 1924-1953). In all his works, he was
able to demonstrate his beautiful artistic skill, and
some of his best watercolours have recently been
published in their original size (Séguy 2004).
***
In France, up to the second half of the twentieth
century, all distinguished and other dipterists were
Séguy’s students, or Séguy’s students’ students. All
the entomo-parasitologists who were trained at the
Pasteur Institute, also attended Séguy’s lectures at
the Muséum. In the meantime, a new scientific institution was created in 1943: the Office de la
recherche scientifique coloniale (ORSC), which was
aimed at improving knowledge of every aspect of
French colonies, including Medical Entomology. In
1949, the ORSC became the Office de la recherche
scientifique d’Outre-mer (ORSOM), later the Office
de la recherche scientifique et technique d’Outremer (ORSTOM), and yet later the Institut de
recherche pour le développement (IRD). This institution, under its successive names, administered a
series of local facilities in the French colonies (and
later in new independent states), which cooperated
with local Pasteur Institutes wherever both were
present. Entomologists working in these various
units published a long series of articles and monographs which form a significant contribution to the
studies of Diptera3.
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Scientific collections, Tropical Medicine and the development of
Entomology in Brazil: the contribution of Instituto Oswaldo Cruz
M. Romero Sá
Casa de Oswaldo Cruz, Fiocruz, Rio de Janeiro, Brazil.
Abstract. The entomological collection of the Institute Oswaldo Cruz is one of the most representative of
neotropical insects, comprising a diverse variety of specimens of distinct taxonomic groups, including
those not linked to research in tropical medicine. The present work retraces the history of the collection
and reports on its main actors and their professional relationships, emphasizing the peculiarity of such an
important collection still being housed in a medical research institution.
Key words: scientific collection, entomology, Manguinhos, Brazil, tropical medicine.
The emergence and tradition of entomological studies
at the Institute Oswaldo Cruz (also known as Instituto de Manguinhos) developed in parallel with the
history of the institution. Soon after its foundation in
1900, insect taxonomy had become a major part of
the early work carried out at the Institution, especially on Diptera – a group discovered by the end of the
19th century to include vectors of pathogenic parasites. Yellow fever and malaria were of great concern
at the time, and campaigns to eradicate such diseases
in Brazil led to the search for other arthropods as
potential vectors of these and other tropical diseases.
It was during one of such expeditions that hemiptera
were found to be vectors of vertebrate pathogens
(e.g. Chagas’ disease), a discovery that stimulated taxonomic studies on Hemiptera at the Manguinhos. In
subsequent years several other insect groups became
of interest to medical science, which ultimately led to
the formation of entomological research collections.
In time, entomology became a major field of research
in itself at the Manguinhos, being no longer being
restricted to species of medical interest. Generations
of researchers and collectors contributed to the
growth of the Institute’s entomological collections.
The holdings allowed entomologist Ângelo Moreira
da Costa Lima (1887-1964) to produce one of the
most important piece of work ever published on
South American entomology: the 12 volume “Os
Insetos do Brasil”.
The present paper will place the development of
entomological studies in Brazil in perspective, focusing on the work carried out at the Institute Oswaldo Cruz and the formation of its insect collection,
uniquely a large reference collection created in an
institution dedicated to medical research. Today the
entomological collection comprises some 4,000,000
specimens, many of which are type specimens, and
includes material representing nearly all orders of
insects. This essay will explore the beginnings of
Correspondence: Magali Romero Sá, Casa de Oswaldo Cruz,
Fiocruz, Av. Brasil, Manguinhos 21045-900, Rio de Janeiro,
Brazil, e-mail: magali@fiocruz.br
entomological study at the Institute in the two first
decades of the 20th century, a period when research
was linked exclusively to Medical and Veterinary
Entomology. Regarded nationally as a pioneer in this
field of research, the Manguinhos soon consolidated
its reputation as a centre for entomological studies.
I then describe the transformation of the Manguinhos into a centre of General Entomology, where
research could be developed without being restricted to insects of any medical interest. From the
1930s to the 1960s the collections grew exponentially, including many different groups of insects.
This period, which may be considered the golden
age of entomological research at the Manguinhos,
led to the formation of one of most important insect
reference collections outside the natural history
museums in Brazil. It was then that entomologist
Costa Lima initiated the publication of his monumental work on Brazilian insects, and helminthologist Lauro Travassos (1890-1970) began to publish
on his collection of butterflies, simultaneously transforming his helminthological laboratory into an
open space for the study of Insect Taxonomy. Finally, I focus on the period of declining interest in the
institutional entomological collection, which
occurred during the 1960s and 1970s, when political interventions affected the institution as a whole.
Recognition of the scientific value of the collections
was restored in the 1990s, when research into geographically traceable samples of organisms gained
new momentum as a result of studies on global biodiversity and Molecular Biology.
The origin of the collections: Medical and Veterinary
Entomology at the Institute Oswaldo Cruz
Specimen number one in the entomological collection of the Institute Oswaldo Cruz is an Anopheles
collected in a malarial habitat in the city of Rio de
Janeiro, described in 1901 by Oswaldo Cruz (Cruz,
1901). Once the role of mosquitoes in transmitting
malaria was definitely established by Giovanni Battista Grassi and his collaborators, and by Ronald
Ross in 1898, the search for possible mosquito vec-
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M. Romero Sá - The entomological collection of the Institute Oswaldo Cruz
tors attracted the attention of physicians around the
world, including in Brazil. Oswaldo Cruz (18721917), Francisco Fajardo (1864-1906), and Emilio
Goeldi (1859-1917) were among those who were
attracted to the study of malaria and its vectors.
Cruz dedicated the species he described in 1901 to
Adolpho Lutz (1855-1940), then director of the
Instituto Bacteriológico de São Paulo and a pioneer
in medical entomology in Brazil (Benchimol & Sá
2006a)1. Regarded by Cruz as a savant and a master, Lutz had worked on Diptera since the late 19th
century, assembling a collection of this group of
insects. His knowledge of Diptera quickly became
known when he became part of a global network
formed to supply mosquito specimens to the British
Museum for the production of a catalogue of the
mosquitoes of the world. Lutz not only sent specimens to the British Museum but also began to communicate directly with Frederick Theobald (18681930), an entomologist at the South-Eastern Agricultural College, Kent, charged with preparing the
monograph for the British Museum. Lutz became
Theobald’s major collaborator in taxonomic matters
during the six years they exchanged correspondence.
The British even adopted a new taxonomic schema
of Culicidae proposed by Lutz in 1904, which he
regarded by far the best general classification of the
Culicidae yet proposed (Benchimol & Sá, 2005;
2006b). Lutz’s new schema was published as part of
the first doctoral thesis on mosquito taxonomy ever
undertaken in Brazil, for which he acted as supervisor. The thesis Mosquitos do Brasil was written by
the physician Celestino Bourroul (1880-1858)2 and
its excellence won him a prize journey to Europe
(Benchimol & Sá, 2006b). Lutz’ interest in mosquitoes and other insects lead him to put together a significant collection from different parts of his native
country, including specimens obtained personally
and through a network of collectors3. Among the
latter were servants and researchers from the Instituto Bacteriológico, personal friends such as Oswaldo Cruz, Joseph Foeterlle, Carlos Moreira, and
Adolpho Lindenberg, and German immigrants functioning as professional collectors from southern
Brazil, some of whom worked for Lutz for several
generations4. Lutz took the grater part of his collection of Diptera with him to the Instituto Oswaldo
For Goeldi’s work on mosquitoes, see Sanjad (2003).
Born in São Paulo in 1880, Bourroul studied medicine in
Bahia, where he presented his doctoral thesis. He later took
over the position of Émile Brumpt as Professor of Parasitology at the Faculdade de Medicina of São Paulo (Benchimol &
Sá, 2006b). For the relationship between Brumpt and Bourroul, see Opinel & Gachelin (2005).
3
In his work on Tabanidae published in 1905, Lutz listed
all the collaborators who had donated specimens for his
Tabanidae collection. See Lutz, 1905 in: Benchimol & Sá,
(2005: 77).
4
For Lutz correspondence with collectors, see the Museu
Nacional Archives, Fundo Adolpho Lutz. See also http://www.
bvsalutz.coc.fiocruz.br/html/pt/home.html
1
2
Cruz when he moved from the Instituto Bacteriológico, São Paulo, to the Manguinhos in 19085.
During the thirty years Lutz spent at the Manguinhos, his insect collection expanded considerably,
incorporating new groups. The collection was officially incorporated to the Manguinhos’ scientific
holdings after his death in 1940.
At the beginning of the 20th century, entomological studies in Brazil were still in their infancy. Few
contributions had been published on insects during
the19th century, all relating to agricultural subjects or
insect biology, such as those published by Fritz
Müller (1822-1897). A German naturalist who had
migrated to southern Brazil in 1850, Müller became
famous after publication of his work Für Darwin
(1964), in which he argued in favour of Charles Darwin’s theory of natural selection 6. Müller was hired in
1876 by the Museu Nacional, the Brazilian national
museum of natural history, as a travelling naturalist.
Although he never left the southern state of Santa
Catarina, where he lived, Müller became one of the
most productive collectors and contributors to the
Museum’s journal, the Archivos do Museu Nacional 7.
He published ten articles on entomological subjects
in the Archivos from 1877 to 1879, including contributions on the natural history of the butterflies Tricoptera and Blepharoceridae (Gualtieri, 2003: 45-96;
Papavero, 2006:15). Another article on entomology,
which appeared in the Museum’s journal in 1878,
was by Nicolau Moreira and described butterfly
metamorphosis. Other significant names were
Rodolpho von Ihering, director of the Museu Paulista
in São Paulo, who published five articles in 1898-99
on Agricultural Entomology, and Emilio Goeldi,
director of the Museu Paraense, later Museu Emílio
Goeldi, in the Amazon region, who published an article on insects linked to agriculture and another on
Coleoptera (Costa Lima, 1936)8.
The systematic studies of insect vectors initiated
around the beginning of the 20th century gave rise
to a new age in Brazilian Entomology, which influenced the scientific community and stimulated taxonomic research in the country. After the pioneering
5 Lutz insect collection left at the Instituto Bacteriológico de
São Paulo was later donated to the Instituto Butantan. For a
list of specimens deposited at Instituto Butantan see the
Adolpho Lutz Virtual Library (www.bvsalutz.coc.fiocruz.br).
6 In the book, Müller shows the development of Crustacea,
calling attention for larvae morphology, sexual dimorphism
and polymorphism as supports to Darwin’s theory of natural
selection. For detailed information on the life and work of
Fritz Müller, see Papavero (1971) and West (2003).
7 The Archivos, created in 1876, was the main vehicle for
publication of studies in natural history. With the creation of
new regional museums in the late 19th century, such as the
Museu Paulista in São Paulo and the Museu Goeldi in Pará,
new journals were created, offering new opportunities for the
publication of works on natural history (Lopes, 1997).
8 Besides those cited above, five other papers or notes on
insects were published during the 19th century, either in journals other than the Archivos or in independent publications
(Costa Lima, 1936).
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work of Oswaldo Cruz on Anopheles in 1901, Lutz,
then Director of the Instituto Bacteriológico de São
Paulo, published an overview of bloodsucking
insects in 1903. Two years later, in 1905, he started
a series of publications on the mosquitoes of Brazil
in the journal Imprensa Medica de São Paulo, in
which he described an impressive number of new
species based on the material gathered over many
years. When Lutz began publishing on Diptera, he
mentioned that his collection included about 200
species of Brazilian haematophagous Diptera, which
had been collected mainly in the States of São Paulo
and Rio de Janeiro. During this period Lutz began
to publish on yet another group of Diptera in which
he had special interest: the Tabanidae. This work
was published in the Revista da Sociedade Scientífica de São Paulo in 1905. Before he moved to the
Manguinhos, he published two more papers on the
Tabanidae. At the Manguinhos he continued to
work on this group of Diptera, and published his
first paper with a collaborator, Arthur Neiva (18801943), a young physician who would become the
Institute’s most active entomologist.
Lutz arrived at the Manguinhos in 1908, which was
also the year in which Neiva and Carlos Chagas
(1879-1934) were appointed assistants there, officially becoming part of the staff. The two young physicians soon became Cruz’s closest collaborators in
malaria research and in the formation of the institutional entomological collection. Born in Salvador,
state of Bahia, in 1880, Arthur Neiva studied medicine in Rio de Janeiro at the Faculdade de Medicina,
where he defended his thesis on the applications of
the anesthetic stovaine in 1905. Neiva was recruited
to the campaign led by Oswaldo Cruz to combat yellow fever, having left the “Serviço de Profilaxia da
Febre Amarela”, after defending his thesis. Recommended to Oswaldo Cruz by his former professor
Antonio Pacheco Leão, Neiva began work on malaria vectors at the Instituto Oswaldo Cruz in 1906.
Cruz was at this time deeply involved with the creation of an institutional collection of Diptera. As a
specialist on the subject, Lutz became his principal
adviser, helping him with taxonomic problems and
exchanging specimens. In a letter to Lutz in August
1906, Cruz commented: “we carry on with the study
of mosquitoes, trying to increase our collections and
I ask you not to abandon us … we are preparing a
great nursery for breeding and studying the life habits
of live mosquitoes, as well as the transmission of
malaria by the Brazilian anophelines”9. In that year,
Chagas published at an article on the prophylaxis of
malaria in the journal Brazil-Medico, in which he
focused on the known mosquito vectors (Chagas,
1906). Simultaneously, Neiva, then a newcomer to
the institution, published a description of a new
species of Culicidae (Myzomyia tibiamaculata) col9 See Biblioteca Virtual Adolpho Lutz, Correspondência,
Oswaldo Cruz, 31/08/1906. http://www.bvsalutz.coc.fio
cruz.br/html/pt/home.html
189
lected by Chagas in Minas Gerais. Also in 1906 Cruz
published a new genus and species (Chagasia neivae)
based on the material obtained by Chagas. In 1907,
Cruz published yet another two new species of
Anopheles recovered from donated material originating in São Paulo. Chagas, in turn, published two new
species of Anopheles from Minas Gerais and a new
Taeniorhynchus justamansonia (Chagas, 1907a, b,
and c). Through these publications, and the creation
of the entomological collection, the Manguinhos
started to build expertise in a new field of medicallyrelated research – insect taxonomy.
Unlike Neiva, Carlos Chagas had taken malaria as
a major research focus from his early days as a medical student. He had worked with his professor,
Francisco Fajardo, at his laboratory at the Santa
Casa de Misericórdia Hospital, where he became
acquainted with the haematology and parasite of
malaria. He went to the Manguinhos to undertake
his thesis on this subject, defending it in 1903. After
a short period working at the Jururuba Hospital, an
isolation facility for plague victims in the city of
Niterói, he returned to his malaria studies. In 1905
he went to work on malaria prevention in a rural
area of São Paulo where a hydroelectric dam was
being built. The success of this campaign resulted in
various other malaria sanitary commissions involving not only Chagas but also Neiva and Cruz (Cruz,
1910)10. These campaigns, in which horses were
used as bait and collecting activities were intense,
provided unique opportunities for collecting many
groups of Diptera, so enriching the institutional collection.
Neiva reported on these activities when working in
Xerém, on the outskirt of Rio de Janeiro, in 1907. He
and Chagas had been commissioned to work on
malaria prevention during a project to harness the
waters of the rivers Xerém and Mantiqueira to supply piped water to the city of Rio de Janeiro. Chagas
was soon dispatched to work on malaria control in a
rural area of Minas Gerais, where a railway line was
being built. Neiva remained in Xerém a further 11
months, which according to his words, “… represented a standing point for studies on malaria; all the
elements being found in such vast field of observation
and experience” (Neiva, 1940: 172). Here Neiva was
able to develop studies of the biology and taxonomy
of vector mosquitoes, the results of which were published as three authoritative works on Brazilian
anophelines (Neiva, 1906, 1908, 1909).
While in Minas Gerais, Chagas made an astonishing new discovery: a hemipteran insect which he
identified as vector of a new human trypanosome
(later to be named the Chagas’ disease, the greatest
discovery ever made at the institution)11.
10 Cruz went to the Amazon region in 1910, to fight malaria among constructors of the Madeira-Mamoré railway line.
11 For Chagas’s discovery and its impact on tropical medicine, see Kropf (2006); Sá (2005); Delaporte (2003); and
Benchimol & Teixeira (1994), among others.
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The Institute also received donations of insects of
many groups from different parts of the country.
Papers on the taxonomy of mosquitoes began to be
published based on the newly formed Manguinhos
collection.
Although Chagas began his scientific life publishing on Insect Taxonomy, it was Neiva who became,
in Cruz’s words, the “institution arthropodist” 12.
Neiva developed a close interaction with Cruz on
entomological matters. This resulted in a productive
partnership, and the two jointly supervised the first
doctoral thesis written at the Manguinhos, by medical student Antônio G. Peryassú, on the taxonomy
of mosquitoes. As Lutz had done earlier in the thesis of his student Bourroul, Neiva described three
new species of Culicidae in an appendix to
Peryassú’s thesis (Myzorynchella gilesi; Sabethes
purpureus and Megarhinus fluminensis)13.
The intense exchange of information between
Cruz and Lutz regarding entomological systematic
matters has been demonstrated by Benchimol & Sá
(2006a: 355-360), who argue that Lutz was then
considered the foremost authority on insect taxonomy. They also emphasize the strength of Lutz’s influence on the new lines of research being initiated in
the Institute, such as the taxonomic work on ticks
by the young physician Henrique de Beaurepaire
Rohan Aragão. Beginning with an interest in species
involved in the transmission of diseases to man and
cattle, a collection was initiated and in less than
three years the first thesis on Brazilian ticks (supervised by Aragão) had been completed by Carlos
Rohr at the Medical Faculty of Rio de Janeiro. In
1911 Aragão began to publish his own work on the
collection he managed (Aragão, 1911). Although he
successfully navigated such different areas of knowledge as Protozoology and Virology, he never completely abandoned the study of ticks. He kept the
collection active by exchanging specimens and information with several important scientific institutions
and researchers over many years, gaining international recognition. The tick collection remains today
one of the most important held by the Institute.
When Lutz arrived at the Manguinhos in 1908,
the work of preparing and studying entomological
specimens was already established as an institutional routine. Specific books on Entomology and other
zoological groups of the Neotropic Region were
being acquired. Neiva was responsible for organizing the Institution’s library and supplied it with all
the main international journals in the areas of interest to the Manguinhos researchers. As recalled by
12
Letter of Oswaldo Cruz to Lutz in 1908. Fundo Oswaldo Cruz, Correspondência. Departamento de Arquivo e Documentação da Casa de Oswaldo Cruz / Fiocruz.
13
Peryassú had the same trajectory as Neiva. Born in Pará,
Peryassú started his medical studies at the Medical Faculty of
Bahia, later moving to Rio de Janeiro to conclude his studies.
The thesis was published in 1921 in the Archivos do Museu
Nacional. See Benchimol & Sá (2006b: 356-7).
Pinto (1932:7): “such is one of the most brilliant
and less known of Neiva’s accomplishments”.
Neiva was soon to become Lutz’s first co-worker,
jointly publishing six works on Diptera, more specifically on the Muscidae, Phlebotominae, Megarhininae and Hippoboscidae (the latter also in co-authorship with A. Costa Lima) (Benchimol & Sá, 2006).
Their first works were published immediately on
Lutz arrived at the Manguinhos. One was a study
based on the Institute’s collection of Tabanidae, created by Neiva during his work in Xerém in 1907.
Cruz maintained a special regard for the young
Neiva, and during his visit to America expressed the
wish to send him to the US for training in entomology. Cruz was much impressed by the American
institutions he visited, especially the US National
Museum and its resident entomologists Leland O.
Howard, Harrison G. Dyar and Frederick Knab.
Writing to Neiva, he stressed: “what admirable people they are! … I visited Howard and he appeared
well acquainted with our work and introduced me
to his collaborators, our well known Dyar and
Knab. They are working on a beautiful monograph
on the Culicidae based on the study of the larvae.
They showed me their drawings: wonderful! They
will pull the rug from under Theobald. I promised
them the most complete collection possible of our
mosquitoes, most of which they do not know. I
expressed the desire to send an assistant up here to
study entomology with them, and they were most
enthusiastic about the idea”14.
Neiva went to the United States to specialize in
entomology in 1910. He was the only Manguinhos’
researcher of the time to be trained in Insect Taxonomy at a natural history museum15. He got the
most out of his stay and, to complete his education,
went to Europe with a recommendation letter from
Leland Howard to examine South American collections kept in Europeans natural history museums.
During his visit, one specific insect group commanded his attention: the Reduviidae, more specifically the Triatominae, vectors of Chagas’ disease.
Studying the European collections which included
material described by the old masters Laporte,
Latreille, Kulg, Stal, Berg and others was central to
taxonomic work on Reduviidae, and Neiva was full
aware of this (Brenner & Stoka, 1988)16.
14
Letter from Cruz to Neiva, 18.07.1907. Fundação Getúlio
Vargas, Centro de Documentação de História Contemporânea
do Brasil (CPDOC), Arquivo Arthur Neiva, ANc 03.05.25.
Theobald’s catalogue was the main reference for Manguinhos
work related to mosquito taxonomy. Howard, Dyar and Knab
produced a work in four volumes on mosquitoes from North
and Central America, and the West Indies, which reviewed
Theobald’s taxonomy criteria. See Benchimol & Sá (2006).
15
At that time Manguinhos researchers traveled abroad for
specialization. Lutz had already visited European natural history museums to examine types in 1905 (see Benchimol & Sá,
2005).
16
During the XIX Century, Brazil was visited by several foreign naturalists who practically traveled throughout most of
the Brazilian territory to collect specimens for natural
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It was Neiva who identified the Triatoma sent by
Chagas from Lassance, and he later published a
study of the insect’s biology. In 1925, he recalled the
day of the identification: “I recall the early days of
the research to identify the insect received from Lassance after Oswaldo Cruz urgently asked for it to be
done. I still remember those days with “saudade”,
the hope of overcoming the obstacles, without assistance, collections to consult, almost without books
– and prompted by an unforgettable master who has
asked you to identify an insect with the smiling
threat of sending it to a foreign expert for identification” (Neiva in Pinto Dias, 2004: 3).
Neiva soon became a specialist on the Reduviidae,
and built a comprehensive collection by his own field
work, and exchange and donation from different parts
of the country and elsewhere in Latin America.
Besides his work on Taxonomy, Neiva established a
methodology to maintain and breed insects in the laboratory, which he developed to observe their development and study the biology of the different species.
Based on the institutional hemiptera collections.
and on studies he had carried out when in the US
and Europe, Neiva (1914) wrote his thesis on the
systematics of the Triatominae. As stressed by Fonseca (1974): “… Neiva`s study of the types of Triatominae when visiting overseas collections was
fundamental to his work in Brazil, and led him to
establish the basis of systematic study of this group
of insects which are of such great medical and economic importance”.
Neiva achieved growing reknown through his scientific work. In 1915 he was invited by the Argentinean government to organize the Zoological and
Parasitological Section at the Instituto Bacteriológico
de Buenos Aires. Back in Brazil, he went to São Paulo
in the following year to assume the direction of the
Serviço Sanitário do Estado de São Paulo from 1916
to 1918. In 1919 he was invited to go to Japan where
he lectures to various conferences. When back in
Brazil in 1920 he resumed his research on the Reduviidae at the Instituto Oswaldo Cruz. It was during
that time that he attracted his first disciple, César Pinto, who was followed in the 1930s by Herman Lent.
These two represent the second and third generation
of Manguinhos entomologists17.
Divulging entomological knowledge: the creation of
Manguinhos’ technical courses and its scientific journal
In 1908, when the Manguinhos was officially recognized as a scientific institution devoted to the study
history museums. Such material was deposited at different
institutions around Europe. For details on the history of
Diptera collecting in Brazil, see Papavero (1971).
17 In 1923 Neiva was again absent from the Instituto Oswaldo Cruz, assuming the direction of the Museu Nacional, a
position he kept until 1927. He became involved in many scientific and political issues, only returning finally to the Manguinhos in 1937, where he stayed until his death in 1943. See
Borgmeir (1940) and Silva (2006).
191
of infections and parasitological diseases of humans,
animals and plants, its new charter required the
introduction of teaching and the creation of a scientific journal as institutional prerogatives18.
The Manguinhos began to offer, free of charge, a
course on Microbiology and Medical Zoology which
was attended by medical students, veterinarians and
pharmacists. The classes were both theoretical and
practical, and were provided by the Manguinhos’
staff 19. Entomology was part of the curriculum, and
Neiva the senior lecturer on the subject. Adolpho
Lutz was in charge of teaching Medical Zoology. In
1913, the entomology programme included studies
of Diptera, Hemiptera and Siphonaptera (Fonseca
Filho, 1974). In the 1919 charter, practical activities
were emphasized and the course became known as
“Applied Course”. The institutional scientific collections were essential to practical training, and were
still growing in size and importance at this time. In
1931 the entomological course incorporated the
Arachnidae, and had its scope greatly extended.
From their creation, the Manguinhos courses were
always the most prestigious in Brazil, educating generations of specialists in public health and tropical
diseases, especially in Parasitology and Medical
Entomology. Their relevance was recognized
throughout Latin America, being attended by students from Argentina, Bolivia, Colombia, Costa
Rica, Ecuador, Nicaragua, Paraguay and Peru (Fonseca Filho, 1974: 134).
An important initiative for the publication of the
work produced at the Manguinhos was the creation
of the journal Memórias do Instituto Oswaldo Cruz,
which first appeared in 1909. Before this date,
works on the taxonomy of vector insects were published in medical journals such as the Brazil-Medico,
the Revista Medico-Cirurgica do Brasil and the
Imprensa Medica de São Paulo. Specialized journals
devoted solely to Entomology were only founded in
Brazil in the 1930s. It was on the initiative of a
Franciscan monk of German origin, Father Thomaz
Borgmeier, that the Revista de Entomologia was created in 1931 and circulated until 195120. During the
20 years of its circulation the journal was directed
and edited by Borgmeier himself, and survived
18 The Manguinhos was created as the Instituto Soroterápico Federal, but in December 1907 its name was changed to
Instituto de Patologia Experimental. Soon after, in March
1908, the name was again changed to Instituto Oswaldo Cruz.
For details of this institutional history, see Aragão (1950),
Fonseca Filho (1974), Stepan, (1976) and Benchimol (1990),
among others. For additional information on the Manguinhos
Application Course, see Pinto (1937).
19 From its creation in 1900, the Manguinhos accepted students for laboratory practice without any formal lessons, giving instruction in sterilization techniques, instrumental and
equipment manipulation and so on. The official recognition of
these courses happened in 1908. With time, a more rigid evaluation system was adopted, including periodical evaluations
and examinations. Schwartzman (1984), Pinto 1937), Benchimol (1990).
20 For Borgmeir biographical data, see Kempf (1976).
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through private and institutional subscriptions and
advertisements. In 1957, Borgmeier invested again
in yet another entomological journal, the Studia
Entomologica, which he linked to the publishing
house of his religious order: the Editora Vozes. This
latter journal survived until 1971, a few years before
Borgmeier`s death in 1975. Another entomological
journal appeared in 1948 as the official journal of
the Sociedade Brasileira de Entomologia (Brazilian
Entomological Society), which had been found in
1937. Its Boletim da Sociedade Brasileira de Entomologia, later Revista de Entomologia, is still in circulation today as the main vehicle for the publication of entomological research in Brazil (Rangel,
2006: 177-181).
Other journals in which entomological studies
were extensively published were those linked to
Agricultural Entomology, a field which incorporated
systematic studies and was institutionalized at the
beginning of 20th century 21 . Journals such A
Lavoura, O Campo and Chácara and Quintais were
among the most popular devoted to agricultural
subjects22.
The first number of the journal Memórias do
Instituto Oswaldo Cruz included three papers on
insect taxonomy, all of which were based on material from the institutional collection. One, by Neiva,
was on the Anopheles mosquitoes and their relation
to malaria; the other two, by Lutz and Neiva, were
on the taxonomy of the Tabanidae. The second number of the journal included an authoritative article
by Lutz on the Simulidae. The following year, in
1910, Arthur Neiva published his studies on the
biology of the hemipteran bug, later proved to be
the vector of the American trypanosomiasis, in the
Memórias (Neiva, 1909; Lutz & Neiva, 1909a, b;
Neiva, 1910)23. Recognition of the relevance of the
collections was confirmed by the publication of a
pamphlet in 1909, which listed the scientific holdings of the Institute. Not surprisingly, the best represented insect group in the collections was the
Diptera, with 95 species of mosquitoes and 145 of
flies. The Aragão’s collection of Ixodidae was the
reported to include 40 species.
Another initiative lay in the institutionalisation of
a complementary technical service for illustration,
photography and cartography. Such services were
essential to the taxonomic work developed with the
21 In 1910 the first laboratory dedicated to the study of agricultural entomology was created at the Museu Nacional, in
Rio de Janeiro, under the coordination of zoologist Carlos
Moreira. In 1921 it was transferred to the Instituto Biológico
de Defesa Agrícola, also in Rio de Janeiro (Rangel, ibidem).
See also Moreira (1929).
22 In a thesis on Ângelo Moreira da Costa Lima and the
birth of systematic studies on agricultural entomology in
Brazil, Rangel (2006) gives an account on the scientific journals devoted to natural history and agriculture entomology in
Brazil.
23 For Lutz’s entomological works, see Benchimol & Sá
(2006).
institutional insect collections, and developed a
close association between the technicians and the
researchers. Lutz was among those who benefited
greatly from the skilled technical service provided
by the illustrators. His entomological works were all
beautifully illustrated, especially his work on the
Tabanidae, which was accompanied by drawings
skillfully produced by Manuel de Castro e Silva.
Appointed in 1908, Castro e Silva was the first and
one of the best illustrators at the Institute, where he
worked until his death in 1934. He was able perfectly to blend technical and artistic abilities, a rare
accomplishment in an illustrator expected to reproduce in perfect detail an organism (or part of it)
selected by the researcher. A single deviation from
the natural model could destroy the scientific value
of the work24. Another skilled illustrator of insects
was the German Carl Rudolph Fischer (18861955), who was hired as illustrator in 1912 and left
the Institute in 1915. During the three years spent
there, he worked with Adolpho Lutz, and took
charge of illustrating Lutz’ works on the Megarhininae, Ceratopogonidae and Oestridae. Following the
tradition of some 19th century naturalists, Fischer’s
passion for insects led him to study a group of
Diptera while he was working at the Instituto Biológico de Defesa Agrícola in São Paulo. He became a specialist on the Tylidae, having published 13 articles the
Diptera, Coleoptera and Hymenoptera25.
From pests to wonders of nature:
the transformation of the Manguinhos’ insect
collections and the role played by
Ângelo Moreira da Costa Lima
Until the beginning of the 1920s, entomology at the
Manguinhos followed the lines of research consolidated in the previous decade, which focused on two
great insect groups of medical and veterinary interest: the Diptera and the Hemiptera. At that time, the
Institute was confronting a period of change, following the death of Oswaldo Cruz in February
1917, and Carlos Chagas’ assumption of Cruz’ position as head of the Institute. In 1919 Chagas was
also appointed head of the newly created National
Department of Public Health (NDPH). As stressed
by Kropf (2006: 194-5), Chagas maintained the
links between research, teaching and production,
and public health, as had Oswaldo Cruz. Under
Chagas’ leadership, specific sections were created
and norms established for the organization, conservation and control of scientific collections (Albuquerque, 1997: 2). It was during this period that, in
1927, the entomologist Ângelo Moreira da Costa
24 Oliveira and Conduru (2004) reported a case related to
such problem, which happened in the Institute with a series
of drawings on the Reduviidae, which were discarded because
considered much more artistic than scientific.
25 For biographical data on Fischer, see http://www.
bvsalutz.coc.fiocruz.br
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Lima was invited by Chagas to take charge of the
entomological section of the Institute, a position formerly occupied by Arthur Neiva. Among Costa
Lima’s duties were the entomology lectures on the
Applied Course. Under his direction, the course was
considerably widened in scope, to include general
observations on insects; notions of external and
internal anatomy; insect development and metamorphosis, general classification of insects; international rules of nomenclature; and the study of specific
groups such as lice, hemiptera (Triatominae and
Cimidae), haematophagous Diptera of the suborders Cyclorrhapa (Muscidae and Sarcophagidae),
Siphonaptera and Arachnidae26. It is worth noting
that emphasis was given to taxonomic studies: students learnt all aspects of identification, application
of the international rules of nomenclature, and
insect classification.
The appointment of the entomologist Costa Lima
considerably altered the profile of the Manguinhos’
entomological collection, turning it into one of the
most important of Latin America.
Costa Lima was an old associate of the Manguinhos and its researchers. His connection with the
Institute and its staff dated from 1907 when, still a
student, he went to work in the ”Brazilian Prophylaxis Service for Combating Yellow Fever”. As a
qualified doctor, he went to Amazonia with Oswaldo Cruz three years later to combat yellow fever in
the city of Pará. There he was charged with identifying the focus of the disease in the cities of Santarém and Óbidos. Study of the insect vectors
improved his knowledge of the taxonomy and biology of insects, especially of the Diptera. On occasion he also experimented with, and put into practice, biological methods of fighting mosquitoes.
Back in Rio in 1913, Costa Lima went to the Manguinhos to re-establish contact with Cruz, who
assigned him to Lutz’ laboratory, where he began
work on other groups of insects besides the
Diptera27. He worked there for a year, when he was
appointed to teach Agricultural Entomology at the
School of Agriculture and Veterinary Medicine,
making him the first professor of Agricultural Entomology in Brazil28. Costa Lima eagerly embraced his
new responsibilities. He began to build collections
26 See Fundo Instituto Oswaldo Cruz, Série Departamento
de Ensino e Cursos, Curso de Higiene e Saúde Pública, Maço
2, 1930. Departamento de Arquivo e Documentação da Casa
de Oswaldo Cruz / Fiocruz.
27 At the Manguinhos, Costa Lima developed three projects,
one in collaboration with Lutz and Neiva and two others
based on material collected by Cruz and Lutz and not linked
to either medical or veterinary taxonomy. These papers deal
with genuine insect taxonomy, indicating their interest in
assessing the diversity of local entomofauna. After leaving the
institution, he published an additional work with Lutz on
fruit-flies (Lutz & Costa Lima, 1918). For Lutz works, see
Benchimol & Sá (2006).
28 For an extensive account on Costa Lima’s life and work,
see Rangel (2006) and Bloch (1968).
193
in the School with the help of his students, professional collectors and donators. He started to publish
extensively in the new journals created to disseminate agricultural matters. In a few years he became
the foremost authority in Brazil on insects of agricultural interest, and was invariably called on to
resolve all problems relating to this field.
Invited to return to the Instituto Oswaldo Cruz,
Costa Lima kept his position at the School of Agriculture. Resuming work at the Manguinhos, Costa
Lima returned to his studies of insects of medical
and veterinary relevance. His first official report
(1927) reflects the intense work developed in the
Manguinhos entomological section under his supervision, including the reorganization of the entomological collection and the incorporation of new specimens; public consultations on systematics and general and applied entomology; and the organization
of an insectarium. Research work reported included
investigations on Anopheles larvae, a Coleopteran
ectoparasite of murid rats, and a Hymenoptera parasite of Triatoma, among other subjects. In that
same year, Costa Lima published four papers on the
Culicidae as well as articles related to economic
entomology 29.
During 1928-29, Costa Lima was deeply involved
with the outbreak of yellow fever in Rio de Janeiro.
Working with Henrique Aragão, he designed several experiments on Stegomyia and the way it transmits yellow fever. His findings were widely publicized and published in the Manguinhos Journal
(Benchimol, 2001).
Within a few years Costa Lima’s entomological
laboratory at the Manguinhos had become a reference point for taxonomic studies. Along with the
laboratory of helminthologist and part time entomologist Lauro Travassos, the Manguinhos became
a Mecca for young entomologists and specialists
from all over Latin America.
Copious numbers of insect specimens found their
way to Costa Lima via his pupils, field researchers,
and by donation and exchange. A considerable number of specimens were also obtained through the initiative of a wealthy amateur entomologist and collector, who had known Costa Lima since childhood.
The Maecenas Carlos Alberto Seabra, from a traditional middle class family of Portuguese origin, provided Costa Lima regularly with quantities of specimens obtained by professional collectors or by the
purchase of private collections. With the aid of the
Brazilian National Council for Research (CNPq),
Seabra bought the collection belonging to the Czech
Joseph Francisco Zikán, which included about
150,000 specimens of Lepidoptera, Coleoptera and
Hymenoptera, among others, collected mainly in
Brazil’s first national park (i.e. Parque Nacional de
Itatiaia). This he donated to the Instituto Oswaldo
29 See Fundo Instituto Oswaldo Cruz, Relatório do ano de
1927, Seção de Entomologia, Departamento de Arquivo e
Documentação da Casa de Oswaldo Cruz / Fiocruz.
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Cruz in 195230. As a result of such initiatives, the
Manguinhos entomological collection soon became
a reference collection for research into nearly all
entomological groups.
In the mid-1930s, Costa Lima began what was to
become a life-long project of cataloguing Brazilian
insects. For the first volume of the great editorial
project, Os Insetos do Brasil, he republished his
papers on the taxonomy of agricultural insects from
the journal O Campo. In the preface, Costa Lima
explains that “the work is directed to those familiar
with the basic morphology and physiology of insects
… who wish to amplify their knowledge through the
study of each order of insects … or those who wish
to begin the study of Brazilian entomology” (Costa
Lima, 1938, preface). The Insetos do Brasil was
published in 12 volumes, covering 27 insect orders,
between 1938 and 196231.
Although the publication was financed by the
School of Agriculture, it was at the Manguinhos that
Costa Lima undertook this work. There he had the
facilities and support necessary for the taxonomic
and editorial work, and benefited from the large
institutional insect collection (which was growing
exponentially); an exceptional library whose holdings included nearly all the classical works on
Neotropical Entomology; and the technical services
of the resident illustrators and photographers.
The relevance of Costa Lima’s editorial work for
his Insetos do Brasil was recognized by his Manguinhos’ contemporaries, who continued fully to
support him even after he resigned as member of
staff in 1938. In 1937 it was enacted that determined public servants could hold two appointments,
since attention was required full time. Costa Lima
therefore chose to keep the professorship at the
School of Agriculture. The Manguinhos’ directors,
however, permitted him to keep his laboratory space
and to continue his research work unofficially, providing him with all necessary support to carry on his
life’s project. Costa Lima took advantage of this
opportunity to stay at the Manguinhos until his
death in 1964.
The freedom accorded to Costa Lima to assemble an
entomological collection of wide scientific interest was
similarly extended to the Manguinhos helminthologist
Lauro Travassos. A well known among specialists
with related interests, Travassos was responsible for
the introduction of taxonomic studies on helminths
to the Manguinhos and, indeed, to Brazil. As did
Costa Lima, Travassos had worked in the Service of
Prophylaxis of Yellow Fever, becoming a member of
the institute’s staff in 191332. A part time entomol30 http://www.bvsalutz.coc.fiocruz.(r/html/pt/static/corres
pondencia/joseph.htm
31 Costa Lima had no time to conclude his second volume
on Hymenoptera, as well as the volumes on the Trepsitera and
Diptera (Rangel, 2006).
32 The helminthological collection built by Travassos and his
assistants is today the largest of South America and includes
ogist with a predilection for butterflies, he built an
important collection of Lepidoptera which he kept
in his laboratory for the benefit of the interested.
From 1937 on, Travassos began to publish on the
taxonomy of butterflies, mainly on material from his
own collection. The Lepidoptera collection progressed, and was eventually officially designated as
an activity of his helminthological laboratory33.
Charismatic and an excellent lecturer, Travassos was
a competent teacher on the Manguinhos’ Applied
Course. In the early 1930s, he also lectured on Parasitology at the School of Agronomy and Veterinary
Medicine, attracting many students to the Manguinhos. His enthusiasm for entomological matters led
Travassos always to provide support for those wanting to study the subject: he opened his laboratory to
amateur and professional entomologists alike.
Together with Costa Lima and César Pinto34, Travassos stimulated students to engage in the field of Veterinary Entomology. Specialists in this field of entomology were absorbed by the institution, thus generating new specific collections, such as those of
Fabio Leoni Werneck (1894-1961). A physician and
a pharmacist, Werneck started his entomological
activities at Costa Lima’s laboratory as an unpaid
assistant in 1930. After concluding the Applied
Course (1931-1932), he was appointed as a laboratory “Associate Head” at the Manguinhos in 1933,
dedicating himself to the study of lice of the orders
Mallophaga and Anoplura. Werneck formed one of
the most important collections of this group in
Brazil, having visited museums in Europe and the
US to compare types and study collections35. The
Werneck collection comprises 4,069 slides; the Mallophaga specimens are preserved in 2,823 slides, of
which 817 are type specimens (Cardozo-de-Almeida
et al., 2003).
Costa Lima and Travassos generated a network of
students who circulated between the Manguinhos
and other institutions. Most notorious was the veterinarian Hugo de Souza Lopes (1909-1991) who,
as a Travassos student at the School of Agriculture,
specimens from all Brazilian ecosystems. The collection
presently contains about 36,000 specimens, preserved either
as whole mounts or wet material.
33 See Relatório da Seção de Helmintologia. Fundo Instituto Oswaldo Cruz, Departamento de Arquivo e Documentação
da Casa de Oswaldo Cruz / Fiocruz and Magalhaes Pinto
(2005).
34 César Pinto became a specialist, not only on insects of the
group Hemiptera and Diptera, but also in helminthes and protozoa. Together with Lauro Travassos he started lecturing on
parasitology at the Medical Faculty of São Paulo. As a parasitologist, he published many articles and books on parasites
of medical and veterinary interest (Sá & Lourenço, 2001).
35 Werneck credited his introduction to entomology to Costa Lima, César Pinto and Lauro Travassos. His collection was
formed basically trough his own efforts. He traveled intensively to the Brazil hinterland and abroad, including Africa, to
collect and visit scientific institutions to study lice. Werneck
was the author of two books and more than sixty scientific
articles. Sá & Lourenço (2001).
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195
began working at the Manguinhos as a volunteer in
the Travassos laboratory. There he initiated a collection of Diptera Muscoidea, having published his
first works on the group in 1932 (Lopes, 1932a, b).
After concluding his studies in 1933, he was deputed by Costa Lima to work on Diptera at the Instituto de Biologia Vegetal, in Rio de Janeiro. He
simultaneously assisted Travassos with his lectures
at the Veterinary School. In 1938, he returned to the
Manguinhos to continue his work on flies in Travassos’ laboratory, now as an unpaid researcher. Only
in 1949 did Souza Lopes become a permanent
member of the Manguinhos’ staff. Lopes was a pioneer researcher on the Sarcophagidae, achieving
worldwide recognition. He also put together an
impressive collection of Diptera which was deposited with the Travassos’ laboratory (Oliveira, 1989).
Unlike Costa Lima, who did not appreciate field
work, Travassos enjoyed collecting activities and
promoted a series of field trips to Central Brazil
(Mato Grosso). Such activities gathered specialists
with different interests from different institutions
around the country. In the 1950s, the entomological
collections deposited at the Travassos laboratory
(butterflies, mosquitoes, flies and other less represented groups of insects) were incorporated to the
general entomological collection of the Instituto
Oswaldo Cruz (Oliveira & Messias, 2005).
At present the entomological collection of the
Instituto Oswaldo Cruz contains some four million
specimens with specimens representing nearly every
order of insects.
Herman Lent, a young medical student, joined the
Travassos laboratory in 1932, also as volunteer, having attended the Manguinhos’ technical course.
Inspired by Travassos’ lectures, he decided to study
helminths, having published his first work on nematodes in 1934 (Lent & Freitas, 1934). However,
when Neiva returned to the Manguinhos in 1936
after leave of absence, he co-opted Lent, who succumbed to his charm, to study Hemiptera. Lent
became member of the Manguinhos staff in 1936,
gradually engaging in Entomology rather than
Helminthology. His first works on Hemiptera were
published with Neiva in 1936 (Neiva & Lent, 1936)
and 1939 (Neiva, Pinto & Lent, 1939). This shift in
Lent’s career resulted in his continuing Neiva’s work
after the latter’s death in 1943, and in the considerable enrichment of the Hemipteran collection and
its transformation into a worldwide reference collection36.
The scientific collections of the Instituto Oswaldo
Cruz were formed mainly as a result of surveys carried out from the early 20th century in the context
of sanitary inspections and the search for the causes of endemic and/or epidemic diseases in the environs of large cities (as Rio de Janeiro and São Paulo)
or in the Brazilian hinterland. Instructions for collecting scientific material were then produced. During the sanitary commissions of the Instituto Oswaldo Cruz from about 1912 on, Brazil was extensively surveyed from north to south, and from the
Atlantic to its western borders, researchers having
traveled extensively along the basins of the nation’s
great rivers such as the Amazon, the São Francisco
and the Paraguay-Paraná (see Lima, 1999; Thielen
et al., 1991; Benchimol & Sá, 2007). New insect
vectors were discovered during these expeditions,
opening new lines of research at the Manguinhos, as
was the case with the Hemiptera at the beginning of
the 20th century and with Phlebotomus in the late
1930s. The discovery of visceral leishmaniasis in the
north-eastern part of the country in 1934 resulted in
one of the most thorough surveys of phlebotomines
carried out by Manguinhos’ researchers. Octávio
Mangabeira Filho, a recent medical graduate hired
by the Manguinhos as a specialist technician in
1938, was appointed to study the Phlebotomus collected, thus consolidating the taxonomic studies on
this group in Brazil38.
Although the Manguinhos collections were initiated as a result of medical-related interest, they gradu-
Travassos’ laboratory continued to train entomologists who became professionals not only at the Manguinhos but also at other scientific institutions, such
as the Museu Nacional, the Brazilian national natural history museum. The entomologists based in
Travassos’ laboratory also supervised entomology
students, for example Sebastião de Oliveira (19182005), a veterinarian entered Travassos’ laboratory
in 1939 as a volunteer, while still a student. Supervised by Hugo Souza Lopes, he initiated a study on
flies of the families Clusiidae and Anthomydae. In
1944 he began to study the Chironomidae, becoming a specialist on this group of Diptera. Sebastião
de Oliveira was officially appointed a member of the
Manguinhos staff in 195037.
36
For a bio-bibliography on Lent, see Jurberg & Santos
(2004).
37
Sebastião de Oliveira became curator of the general entomological collection of Instituto Oswaldo Cruz from 1986
until his death in 2005. For biographical data on him, see
Oliveira & Messias. Calaça (2001) gives a sociological analysis of the rotation of personnel between Manguinhos’ laboratories, emphasizing the Travassos’ laboratory.
Conclusion
38
Lutz and Neiva in 1912 had already published on Phlebotomus, and Henrique Aragão in 1922 had experimentally
demonstrated that one of the species described by Lutz and
Neiva was responsible for the transmission of cutaneous
leishmaniasis. However it was only after the discovery by
Evandro Chagas of visceral leishmaniasis in Brazil that systematic study of Phlebotomus became a major interest. Studies on the group were carried out by Octavio Mangabeira Filho (1913-1963) who, between 1941 and 1942 alone, published 12 papers on the taxonomy of Phlebotomus, including
descriptions of 40 new species. See Chagas (1936) and Sherlock (2003).
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ally increased their potential as sources for a better
knowledge of the entomological diversity of the country. The richness of the holdings, incorporating an
immense variety of specimens of different taxonomic
groups collected at distinct localities, makes them an
invaluable source of information on the geographical
distribution of species. Such data are fundamental for
determining the geographical distribution of the vectors, hosts and diseases of animals and humans. Thus
the collections gathered by the first Manguinhos
entomologists are both an essential component of the
taxonomic framework inherent to the species represented, and a vital source of environmental, epidemiological and biogeographical data. Further, they constitute a central part of the institutional historical
memory and of the Brazilian scientific trajectory.
Neglected from late l960s to the mid 1980s, the
collections experienced a period of stagnation and
misuse, during which basic taxonomic studies were
discontinued following political upheaval and new
administrative priorities. This (temporary) disregard
for the potential of the collections as research tools
was enhanced by the introduction of a revolutionary
new line of research into the laboratories: Molecular
Biology. Paradoxically, however, it is precisely
researchers in Molecular Biology who are now turning to the collections for data on the taxonomy and
distribution of parasites and vectors complementary
to their own findings. This recent rediscovery of the
potential use of collections results from the fact that
molecular biology works in a micro dimension, somewhat remote from the spatial dimension of the organism itself. The detection of distinctions between populations of vectors or hosts on the molecular level
demands information on the actual environment
where the organism functions. In this way, all data on
the morphology, biogeography and biology of the
whole organism are essential to substantiate laboratory research and confirm the results of eventual molecular distinctions detected by these methods.
As the world is gradually degraded, ecosystems
destroyed, and populations and species exterminated, the value and interest of examples of natural
organisms will increasingly grow. As far as medical
research is concerned, such examples – besides their
uses as discussed above – may well lead to answers
for otherwise insoluble problems.
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2
Portraits
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Battista Grassi entomologist and the
Roman School of Malariology
E. Capanna
Dipartimento di Biologia Animale e dell’Uomo, Università “La Sapienza”, Roma, Italy, and Centro Linceo interdisciplinare “Beniamino Segre”, Roma, Italy.
Abstract. Grassi’s entomological researches go back to his early years of university study, initially being
concerned with agriculture pests. After working in Heidelberg, Grassi’s interests turned to basic problems
of entomology, such as the evolutionary origin of Miriapoda and Insecta, and termite caste determination.
His first investigations into medical entomology related to the problem of bird malaria, which he studied
only in relation to the hematic parasites. In 1895, Grassi was appointed Professor of Comparative Anatomy at Rome University, and initiated his entomological collaboration with the Roman malariologists, Amico
Bignami, Giuseppe Bastianelli and Ettore Marchiafava. At the end of 1898, they announced, at the session
of the Accademia dei Lincei on December 4th, that a healthy man in a non-malarial zone had contracted
tertian malaria after being bitten by an experimentally infected Anopheles claviger. Following his disappointment at being excluded from the Nobel prize, Grassi devoted his attention to another important insect
related to the transmission of parasitic disease, the sand fly, Phlebotomus papatasii. After World War I
malaria had flared up with renewed vigour, so that the social importance of the disease convinced Grassi
to resume his studies in 1918. The problem he faced in these years was “Anophelism without malaria”
which was to be solved a year after his death by his pupil Falleroni, who demonstrated that there are six
cryptic species of Anopheles of which only four bite humans and transmit malaria. Battista Grassi died on
4 May 1925, working to the end: he was reading the proofs of his last paper, Lezione sulla malaria.
Key words: Battista Grassi, malaria transmission, Anopheles, Phlebotomus.
The entomologist
An Italian zoologist of the XIXth century (Lessona,
1869) wrote “the main qualification for being a perfect naturalist is to be born in the country, where he
will have spent his best years amid natural scenery,
looking at Nature without preconceived ideas”. Battista Grassi was born on 24 March 1854 in Rovellasca, a small country village not far from Como, to
a landowning family of the cultured Lombard middle class. He spent his boyhood years immersed in
nature and this way of life shaped his training as a
“perfect entomologist”. Grassi’s first researches –
carried out while he was still student in the Medical
Faculty of Pavia University – well testify to an inclination for entomology; they deal with agriculture
pests, like certain Hymenoptera (Ichneumon), Lepidoptera (Tinea), and Coleopteran (Meloe) (Grassi
and Parona, 1876, 1877).
Grassi’s interest in the insect world developed an
evolutionary perspective following his studies in Heidelberg, with Carl Gegenbauer (1826-1903), one of
the first Darwinian comparative anatomists. Many of
Correspondence: Ernesto Capanna, Dipartimento di Biologia
Animale e dell’Uomo, Università “La Sapienza”, via A. Borelli 50, 00161 Roma, Italy, Tel +39 06 49918008, Fax +39 06
4457516, e-mail: ernesto.capanna@uniroma1.it
This paper is an extension and a widening of part of a lecture
I held in Barcelona at the Instituto de Estudios de Cataluña
(19 October 2005), published in a short form in International Microbiology (9: 69-74, 2006), titled “Grassi versus Ross:
who solved the riddle of malaria?”.
Grassi’s essays dealing with I Progenitori di Miarpodi e Insetti (i.e, The Ancestors of Myriapoda and
Insecta) (Grassi, 1885, 1886a, 1887a, 1889), as well
as those on the white ant caste (Grassi, 1888a and
b, 1890; Grassi and Sandias 1893) (Fig. 1), belong
to this fundamental entomological research. They
were widely influential internationally, resulting in
the award of the Darwin Medal of the Royal Society
in 1896. In a letter to Grassi, Sir Edwin Ray
Lankester (1847-1929), announced the Society’s
decision, and emphasised that the prize was awarded to “those naturalists who are still in active work
and especially doing work which has important and
direct bearing on Mr. Darwin’s own investigation
and theory” (Lankester 1896 in letter, quoted in
Capanna 1996, p. 37). In the same letter Lankester
added that, before Grassi, the prize had been awarded to Alfred Russell Wallace (1823-1913), Joseph
Dalton Hooker (1817-1911) and Thomas Henry
Huxley (1825-1895). In the opinion of the English
zoologists, therefore, the standing of Battista Grassi,
as naturalist, was equal to that of the three greatest
and most faithful friends of Charles Darwin (letter
dated Oxford, 15 July 1896).
We must not forget, however, that Grassi had a
solid grounding in Pathology. He well knew the
importance of competence in zoology, especially in
its entomological and helminthological aspects, for
solving parasitological problems. In 1888, therefore,
Grassi – now Professor of Zoology and Comparative
Anatomy at the University of Catania – began to
study malaria in birds, in collaboration with the
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E. Capanna - Battista Grassi entomologist
Fig. 1. Preparatory drawings of Grassi’s paper on the cast of the White ants (from the Grassi Archive of the Museum of
Comparative Anatomy).
medical clinician Riccardo Feletti (Grassi and Feletti, 1890). In 1891 they published the monograph
“Ueber die Parasiten der Malaria” in Centralblatte
für Bakteriologie und Parasitenkunde (Grassi and
Feletti, 1891a, 1891b) in which they described the
malarial cycle in different species of birds, such as
the owl, pigeon and sparrow. It was here that we see
his zoological approach to the problem: different
species of birds, belonging to different orders
(Strigiformes, Columbiformes and Passeriformes),
were demonstrated to be parasitized by different
species of protozoans: only Halteridium in pigeons,
but also Proteosoma praecox (Haemoameba) in the
sparrow. This work took place five years before
Ross turned his attention to the study of avian
malaria in India. Yet the Italian scientists did not –
at that time – recognise the true mode of transmission of malaria by means of hematophagous insects.
Grassi and the Roman Malariologists
In 1895, Grassi was appointed Professor of Comparative Anatomy at La Sapienza University in Rome.
Malariological research was a lively and productive field in Italy, especially from the clinical and
public health point of view. But Italian malaria studies had a different perspective from that developed
in France and Great Britain: for the French and
British researchers, malaria was chiefly a “tropical
disease”, a “colonial” problem. In England and in
France, malaria had a very restricted presence in
estuarine marshes areas. For Italian physicians, by
contrast, malaria was a disease endemic to their
own country, unified only a decade earlier, a scourge
that hindered the development of many of its southern regions. Rome, the capital of this young kingdom, was a city in the grip of malaria during the
summer and autumn months. In a famous speech in
Parliament, Giustino Fortunato (1848-1932), a
native of Rionero in Volture, a malarial zone in
southern Italy, warned that until southern Italy was
rid of this incapacitating disease, there could be no
development of the region (Fortunato, 1898). A
map of Italy published in 1882 indicated in red the
areas with widespread malaria, and in yellow areas
where the disease was present (Torelli, 1882). The
red area included vast coastal stretches of Tuscany
(Maremma), Latium (the Roman plain and Pontine
marshes) and Campania. Also at high risk of malaria were the Venetian lagoon and the Po River delta,
the Ionian Coast of Calabria, and the coasts of Sardinia and Sicily. Even more tragically astonishing is
another map published in 1899, produced by professor of hygiene Augusto Celli (Celli, 1899), which
indicated the railway lines where the risk of contracting malaria during a train trip was high!
While for Alphonse Laveran (1845-1922) or
Ronald Ross (1857-1932), malaria was a problem
affecting peoples of remote countries, for Italian
doctors it was thus part of a daily domestic tragedy.
The Grassi Archive that I created in the Department
of Animal and Human Biology (Capanna and Mazzina, 1998) contains the following report among the
clinical records of a malariological investigation
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conducted in Rome and in the Roman countryside
by Grassi and collaborators in 1900. The report is
disturbing in its tone of laconic tragedy; “R. A.,
from Anguillara (Rome), 9 years old. Fever on 2
August […] suffered a severe bout of fever on 22
August, and died of pernicious algid fever on the
night of 23 August in Santo Spirito Hospital”
(Capanna, 1996, p. 12). In Rome at the beginning
of the twentieth century, a nine years old boy died
of malaria within a period of 20 days. This was the
potent stimulus that prompted a group of Roman
doctors to attempt to resolve the problem of malaria. Of these physicians, I will mention only a few
eminent figures.
Ettore Marchiafava (1847-1935), Professor of
Pathological Anatomy, and Celli, taught in the Faculty of Medicine of Rome, where they collaborated
closely in their research on malaria. They were
somewhat perplexed by Laveran’s discovery, since
they believed that the pathogenic factor in malaria
was a Bacillus, B. malariae (Klebs and Tommasi
Crudeli, 1879). However, after careful study, they
agreed with the observations of the Frenchman indicating a protozoan as the malarial parasite. Since
the generic name Oscillaria proposed by Laveran
had been attached to another organism, a green-blue
filamentous alga (Cyanophycaea), in 1885, the two
scientists proposed the name Plasmodium for the
genus (Marchiafava and Celli, 1885).
Amico Bignami (1862-1929), Professor of General
Pathology, was a pupil of Marchiafava, and together
with Giuseppe Bastianelli (1862-1959), clinicianphysician, collaborated closely with Grassi in the
identification of the malarial vector, especially with
regard to the intrahaematic cycle and the clinical and
therapeutic aspects (Bignami, 1896, 1898; Bignami
and Bastianelli, 1898). An important contribution of
Marchiafava and Bignami was the identification of
the two species of Plasmodium, P. falciparum and P.
vivax (Marchiafava and Bignami, 1894).
Camillo Golgi (1843-1926), Professor of General
Pathology at Pavia, was awarded the Nobel prize in
1906 for his contributions to the physiology of the
nervous system, but he was also heavily involved in
research on malaria. Indeed, the agricultural areas
along the Po River had a high risk of malaria, especially where there were extensive rice fields, as in
the countryside around Pavia. Golgi made a notable
contribution to malariology (Golgi, 1886, 1889,
1894) by relating the clinical signs of the fever
episode with the schizogonic phase of the plasmodium and by showing that the so-called tertian and
quartan intermittent fevers are due to the presence
in the blood of two different Plasmodium species (P.
malariae and P. vivax), sometimes present together.
Last, but not least, Battista Grassi (1854-1925)
(Fig. 2).
In Rome, Grassi came into contact with the group
of Roman malariologists, who convinced him of the
validity of the transmission of the Plasmodium via a
hematophagous insect, a hypothesis he had until
203
Fig. 2. Portrait of Battista Grassi forty years old (from the
Grassi Archive of the Museum of Comparative Anatomy).
then he considered doubtful. The problem was to
identify the incriminated insect with certainty, and
the “Roman Malariologist” needed the cooperation
of an entomologist. Grassi began the investigation
with the tools of the entomologist, namely knowledge of the systematics of the group and of the geographical distribution of the species. On the basis of
the epidemiology of malaria and the distribution of
the mosquitoes present in the malarial zones, he
first selected a group of three species suspected of
malarial transmission, Anopheles claviger (synonym
A. maculipennis) and two Culex species (but not
including the common C. pipiens), and he communicated this result to the Lincei Academy on 2 October 1898 (Grassi, 1898a).
On 6 November 1898, Grassi (1898b) announced
to the Lincei Academy that, with Bignami and Bastianelli, he had infected a “volunteer” by exposing him
to the bite of these three mosquito species and on 28
November the formal note on this experiment was
presented (Bastianelli, Bignami and Grassi, 1898).
Suspicion of the two Culex species faded immediately
and the innocent mosquitoes were absolved of the
crime of being vectors of the infection; a further note
by Grassi, Bastianelli and Bignami (1899) was read in
the academic session of 4 December 1898, in which
it was announced that a healthy man in a non-malarial zone had contracted tertian malaria after being bitten by an experimentally infected Anopheles claviger.
The experimental phase ended on 22 December with
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a communication to the Lincei Academy (Grassi, Bignami and Bastianelli, 1899), which described the
whole developmental cycle of the plasmodium in the
body of Anopheles claviger and stated that it corresponded to what Ross had described for Proteosoma
in Culex pipiens in the malarial cycle of birds (Ross,
1897a and b).
The experiment was conducted with exceptional
rigour; the Anopheles mosquitoes were raised in the
laboratory from the larval stage, and starved until they
had bitten a patient who had “semilune” – i.e. Manson’s (1894) crescents – in his blood, the only stage
that could have developed into gametophytes and thus
triggered the gonochoric cycle in the body of the mosquito. A certainly healthy person was then exposed to
the bite of these mosquitoes in a place protected from
the introduction of other mosquito species.
We have archival documents relating to this scrupulous experimental procedure, such as various notes
addressed to Bastianelli who accompanied the mosquitoes, as well as records of expenses, e.g. the payments to the “volunteers” who were bitten, where it
appears that the “semilunari”, those patients with
“crescents” in their blood, were rewarded with more
than two liras, twice the amount given to patients
who had other stages in their blood.
In the first issue of the Annales de l’Institut Pasteur of 1899, there appeared an article dated “Calcutta, 31 December 1898” by Major Ronald Ross
and entitled Du rôle des moustiques dans le paludisme (Ross, 1899). The insect responsible for transmission of the disease was indicated as “moustique
d’une nouvelle espèce”, just as in the note of 1897
it was indicated as a “grey” or “dappled winged”
mosquito, absolutely invalid names for the Linnaean
nomenclature. However, Ross was not a zoologist
and he completely lacked the tools of zoological systematics. Grassi correctly noted in the margin of
Ross’s article “he doesn’t say that it was Anopheles!”. The volume of the Annales de l’Institut Pasteur in our library contains numerous handwritten
notes by Grassi (an angular calligraphy that reveals
a punctilious character) (Fig. 3).
On 4 June 1900, an article by Battista Grassi, entitled Studi di uno zoologo sulla Malaria (Studies on
Malaria by a Zoologist), was published in the Memoirs of the Royal Lincei Academy (Grassi, 1900). It
consisted of 200 large-format pages in which Grassi summarized his four years of research from 1896
to 1899 and underlined the originality of his contribution by defining himself as a zoologist.
In the crucial years of research on the transmission of malaria in Rome (between 1897 and 1898),
the English physician, Dr. Edmonston Charles, visited Grassi’s laboratory in via de Pretis and those of
the other malariologists at the Santo Spirito Hospital. He was greeted without suspicion by the Italian
scientists, who were flattered by the interest of an
English colleague in their studies. Dr. Charles then
reported the information he obtained to Ross. When
the dispute about the priority of the discovery of the
Fig. 3. First page of the article by Ronald Ross “Du rôle des
moustiques dans le paludisme” showing handwritten notes
of Battista Grassi (Library of the Department of Animal and
Human Biology, “La Sapienza” University of Rome).
insect responsible for malarial transmission
emerged, Ross felt obliged to make public the letters
received from Dr. Edmonston. A rare publication by
Ronald Ross, entitled Letters from Rome on the
New Discoveries on Malaria (Ross, 1900) (Fig. 4)
contains several passages from two letters that well
characterize the facts.
In a letter dated 4 November 1898, Dr. Charles
wrote to Ross:
[…] I called on Dr Manson before leaving London
to get the latest news of what progress you had
made in your work, in order to let the Italians know.
They have been working in various directions this
summer, but up till this week without being able to
show any definite results. Bignami has collected
mosquitoes from four very malarious localities.
According to Grassi it would seem there are some
fifty varieties of mosquitoes in Italy. Only six, however, seem to frequent these selected malarial positions.
Besides the mosquitoes, the larvae were also brought
up, and allowed to develop in Rome (Ross, 1900).
Nevertheless, the Roman malariologists began to be
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205
Fig. 4. The pamphlet of Ronald Ross containing the letter sent from Rome by Dr Edmonton Charles (from the Grassi Archive
of the Museum of Comparative Anatomy).
mistrustful towards the Englishman and they did not
tell him the complete truth. In fact, on 4 November,
the suspects were already limited to just three species
and the absolution of the two Culex species had
already been decided. It is noteworthy that Charles
writes “varieties” and not “species” as a good zoologist should have done!
In another letter dated 19 November, Charles
wrote to Ross:
[…] As, doubtless, it would help you to have
named specimens of mosquitoes spoken of by Grassi, I went to his laboratory to try and get him to give
you a few specimens of the different kinds of mosquito. I did this under the impression that he had
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completed his investigations. He told me, however,
they were far from complete, and did not give me
the specimens (Ross, 1900).
Charles’ initial impression was exact. Grassi, Bignami and Bastianelli had by now identified the
malarial vector and on 4 December, only two weeks
after the visit by Dr. Charles, they published their
success in the Reports of the Lincei Academy. The
letter then continues with an interesting sentence:
He (i.e. Grassi) spoke in the highest terms of
praise of your work; he has your first report (i.e. the
note of 18 December 1897 in the British Medical
Journal), and told me to write to try and get your
future reports at an early date for him (Ross, 1900)
On that date, therefore, the relationship between the
two scientists was one of mutual respect, confirmed in
a letter dated Calcutta 5 February 1899 that Ross sent
to Charles, who had sent him the English translation
of the note by Grassi, Bignami and Bastianelli of 22
December 1898. In Ross’s letter, we read:
My dear Dr. Charles, very many thanks for your
last letter with the translation of Grassi, Bignami,
and Bastianelli’s note. This is good indeed. Pray
give them my felicitations.
I thought that the grey mosquito is Culex pipiens,
but was not quite sure. Of course there is a whole
family of allied grey mosquitoes (Ross, 1900) (Fig. 5).
This friendly and collaborative climate continued in
the spring-summer of 1900, when Sir Patrick Manson
(1844-1922) organized a crucial experiment to be
conducted in an Italian malarial zone (Manson,
1900). A small building, in the style of an English
hunting lodge, was designed and built in England and
then assembled in Italy in the Castelfusano pinewood,
on the hunting estate of the kings of Italy near Ostia.
This “mosquito-proof” hut was inhabited by two
“intrepid” doctors, the Italian Luigi Sambon (18651931) and the Englishman G.C. Low, both of the London School of Tropical Medicine, during the period of
the summer-autumn fevers, which is also the period of
maximum reproductive activity of the mosquitoes
(Sambon and Low, 1900, 1901). The “intrepid” doctors, and their servants, remained free of malarial
infection after a stay of three months. The experiment
was also followed by Bastianelli and Grassi, and the
latter sent Manson a telegram dated 13 September
1900: “Assembled in British mosquito proof hut having versified (sic!) [instead of “verified”] perfect
health of experimenters among malaria stricken
inhabitants. I salute Manson who first formulated the
mosquito malaria theory” (quoted in Fantini, 1999).
Nevertheless, at the end of 1900, Ross began a
campaign against the three Italian biologists to
claim the priority of discovery of the mechanism of
transmission of malaria, clearly with the possibility
of winning the Nobel prize in mind. He even put the
originality of Grassi’s research in doubt, maintaining
that Grassi was guided in the identification of the
Fig. 5. Handwritten copy of a letter (dated Calcutta February 5. 1899) sent by R. Ross to E. Charles (from the Grassi Archive
of the Museum of Comparative Anatomy).
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vector by the fact that he had already indicated that
a “grey mosquito with dappled wings” was responsible for the transmission; he also accused Grassi of
fraud on the basis of a wrong dating of the notes
presented to the Lincei Academy, which instead was
precisely certified by the date of presentation in the
Academy’s public session. Grassi reacted vigorously
to these accusations, which in his opinion impugned
his honour as a scientist. The Swedish Academy of
Sciences awarded Sir Ronald Ross the Nobel prize
for Medicine in 1902.
An useless dispute
The dispute between Grassi and Ross about the priority of discovery is usually interpreted as motivated by personal ambition, national pride, the desire
for academic pre-eminence and similar psychological and sociological positions that have very little to
do with science. In this regard, Bynum wrote that
the dispute “[…] is one of the least attractive
episodes in the whole history of malariology”
(Bynum, 1998), and this may be partly true; both
scientists had strong personalities and it was not
easy to find a point of agreement.
Undoubtedly the actual priority concerning the
process of malaria transmission via haematophagous
insect is due to Manson’s (1894) brilliant intuition.
Grassi dedicated his paper Studi di uno Zoologo sulla Malaria (Grassi, 1900) to Patrick Manson. He
placed this dedication on the first page of his monograph: “A Patrick Manson – scopritore del ciclo evolutivo della filaria, geniale iniziatore delle attuali
ricerche sui parassiti malarici” (i.e.: To Patrick Manson discoverer of the life cycle of filarial, clever initiator of the present researches on malaria parasites). Manson did indeed suggest to Ronald Ross an
investigation of the “crescentic and flagellate bodies
in malarial blood”. Although it is true that having
related malarial transmission in birds to a
haematophagous mosquito, as Ross done, albeit not
systematically classified, was by itself a great success
for science, deserving of the Nobel prize, it is equally true that the identification with precision of the
species of nematocerous dipteran that transmitted
malaria in humans must be considered a scientific
success of equal importance. It might have been
objected that, all in all, to have attributed a Linnaean name to the insect was a marginal part of the
problem, but if this might have been justified in the
XIXth century it was no longer so at the threshold
of the XXth century. The current judgment of historians of science attributes – now that the useless
dispute is ended – equal merit to both scientists
(Dobbel, 1925; Corbellini and Merzagora, 1996;
Fantini 1998; Dobson, 1999).
The true cause of the dispute, however, was the
different approach to tackling problems in biological research, in this case, the parasitological cycles.
Grassi’s method was characteristic of zoological
research: systematic, comparative, experimental.
207
The method pursued by Ross was empirical and
intuitive (Fantini, 1999).
Medicine in the 1800s, but also until recent times,
was an empirical science. Not so zoology, which with
the Darwinian revolution tended toward a positivistic
concreteness. For a post-Darwinian zoologist like
Grassi, the question of the species was the focal point
of the process: an animal remained undefined until it
was placed in a context, no longer merely a classificatory context, but also an evolutionary one. The
detailed systematic revision of the European Culicidae, performed by Ficalbi (1896) few years before,
equipped Grassi with a powerful instrument for solving the problem of the insect vector of malaria. For
Grassi, nomenclatural meticulousness was almost an
obsession. In 1899, while preparing to write the article Studies on Malaria by a Zoologist, he needed an
opinion about the nomenclature of the parasitic protozoan of malaria. He turned to the leading expert on
sporozoans, Prof. Raphaël Blanchard (1857-1919),
whose return letter, dated 9 November 1899, is in the
Grassi Archive (Fig. 6). After consulting with various
Fig 6. Letter (dated Paris November, 9) sent by Prof. R.
Blanchard to Grassi where the nomenclature of the different
Plasmodium species has been precisely defined, including
all synonyms (from the Grassi Archive of the Museum of
Comparative Anatomy).
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colleagues, including Alphonse Laveran, Blanchard
provided a scheme of accepted names and different
synonyms for Plasmodium malariae (see Capanna,
1996, p. 39).
The comparative method was the second tool that
guaranteed Grassi’s success. We should remember
that he learned the method from Carl Gegenbaur,
one of the greatest post-Darwinian comparative
anatomists, and that the biological discipline he
taught at the University of Rome was Comparative
Anatomy. The comparative method was widely used
by Grassi not only in the comparison between
species, but also between environments and ecosystems, and again between the species and the environments they inhabited. Thus, in a dialectical
process, he excluded the species that could not be
malarial vectors and unequivocally identified
Anopheles claviger as the sole vector of malaria in
Italy. Grassi wrote: “In medical science, the comparative method must be considered the main route
to arrive at the solution of the problem” (Grassi,
1879). For Grassi, Parasitology was a zoological science; it was the application of Darwinism to Pathology.
Lastly, the experimental method. Progress through
experience was deeply rooted in the tradition of the
Studium of Pavia, the Alma Ticinensis Mater where
Lazzaro Spallanzani (1729-1799), father of Experimental Biology, conducted his fundamental
research. Spallanzani had stated “To experiment is
the work of everyone, to experiment properly is, and
always will be, the work of the few” (Spallanzani,
1782), and Grassi knew how “to experiment prop-
erly”. The meticulousness in designing the experiments involving the experimental infection of mosquitoes born in the laboratory, using patients selected according to the haematic stage of the parasite
and the subsequent biting of certainly healthy
patients in a protected environment, seems to have
been suggested by the cautious and scrupulous
approach of Lazzaro Spallanzani.
The zoologist’s method as used by Grassi had
already brought success in the interpretation of various complex cases of human parasitoses related to
the cycle of helminths. It is sufficient to cite his
analysis of the cycle of Ancylostoma when he was
still a student (Grassi and Parona, 1878), and especially his study of Hymenolepis, a cestode that
ambiguously may have had, or may not have had, an
intermediate host, but which Grassi demonstrated
to be a complex of two species: H. nana, without
the intermediate host, and H. diminuta, which
required two hosts to complete its cycle (Grassi,
1887b).
Anophelism without malaria
Very similar to the case of Hymenolepis, on account
of the nature of its zoological context and the comparative zoological approach with which it was tackled, was the question of “Anophelism without
malaria” (Fantini, 1994). Grassi dealt with this
problem, but did not have time to resolve it completely, even though he deduced aspects related to
the anthropophily and zoophily of several “varieties”
of Anopheles maculipennis.
Fig.7. Preparatory drawings for the lithographic plate of the Grassi memoir “Ricerche sui Flebotomi” (1907) (from the Grassi
Archive of the Museum of Comparative Anatomy).
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Grassi also had his dogma: “There is no malaria
without Anopheles”, but already in 1899, at the
time of the conclusive results concerning Anopheles
claviger, he noticed that there were areas where
Anopheles was abundant but malaria was absent. A
first hypothesis in this regard, expressed in the second edition of his article (Grassi, 1901), was to
relate this phenomenon to the thermophily of the
Anopheles mosquito. In areas with cold nights, the
mosquito did not fly and bite humans, but stayed in
the warmer stalls to bite livestock.
After the disappointment of being omitted from
the Nobel prize, Grassi decided to stop studying
malaria and to devote his attention to Agrarian
Entomology. Important researches on the life cycle
of the vine louse Phylloxera were carried out
between 1907 and 1917, in collaboration with Anna
Foà and a group of good scholars (Grassi, Foà et al.,
1912). However, his passion for Medical Entomology pushed Grassi towards investigating another
insect important in the concern of transmission of
parasitic disease, the sand fly, Phlebotomus papatasii (Grassi, 1907) (Fig. 7).
The social importance of Malariology convinced
Grassi to resume his studies in 1918. Indeed, after
World War I, during which the fight against malaria was abandoned in favour of fratricidal fighting,
malaria had flared up with renewed vigour. Mortality from the disease had rapidly decreased in Italy
since 1898, following the discovery of the vector
and the zooprophylactic activity, and the free distribution of quinine (Coluzzi M., 2004), from 600
deaths per million inhabitants to fewer than 50 in
1915. However, it then increased to 320 per million
in 1919 (Coluzzi A., 1961). Resuming his research,
Grassi turned again to the problem of anophelism
without malaria. He identified three localities with
a typical malarial environment, all infested by
Anopheles maculipennis but not affected by malaria: Orti di Schito near Naples, Massarossa in the
Tuscan Maremma near Lucca (Celli and Gasperini,
1902), and Alberane in the rice fields around Pavia.
In 1921, he demonstrated that “there is certainly a
biological race of Anopheles mosquitoes that does
not bite man” (Grassi, 1921a and b). A year after
his death in 1925, one of his pupils (Falleroni,
1926) showed, on the basis of these observations,
that there are six species of Anopheles in the maculipennis complex, which are indistinguishable
except for their egg morphology (Fig. 8). Of these
six “new” species, Anopheles labranchiae and A.
sacarovi, present in highly malarial zones, mainly
bite man, while the typical form of A. maculipennis,
present at Orti di Schito, only bites animals. The
species present at Massarossa in Maremma, A. messae, mostly bites animals, but sometimes also man.
Therefore, we can see the importance of the precise
systematic identification of the vector for the management of antimalarial zooprophylaxis; it was not
possible to be satisfied with rough identifications
like “grey mosquito” or “dappled winged mosquito”.
209
Fig. 8. Plate of the Falleroni paper “Note sulla Biologia
dell’Anopheles maculipennis”. The eggs of the different
species of the maculipennis complex are shown: 1,
melanoon; 2, messae; 3, labranchiae; 4, maculipennis s.s.;
5 and 6, sacharovi.
Death caught up with Battista Grassi during the
night of 4 May, 1925. He was still in full scientific
activity: he was reading the proofs of Lezione sulla
malaria (i.e. Lecture on Malaria), a rigorous paper
of over 130 pages that was posthumously published
in 1927, as a spiritual testament after near 40 years
devoted to the Medical Entomology.
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Raphaël Blanchard, Parasitology, and the positioning of
Medical Entomology in Paris
M.A. Osborne
Department of History, University of California, Santa Barbara, CA, USA.
Abstract. The histories of medical entomology and parasitology are entwined. Raphaël Blanchard (18571919), Chair of Medical Natural History and Parasitology at the Faculty of Medicine in Paris, organized
the teaching of medical entomology and civilian colonial medicine. He also founded and edited the journal Archives de Parasitologie and started the Institute de Médecine Coloniale where he mentored many
foreign students and researchers. Additionally, Blanchard is important for his scientific internationalism
and medical historical work on the cultural location of parasitology and for training the future professors
of parasitology Jules Guiart, Émile Brumpt, and Charles Joyeux.
Key words: Institute de Médecine coloniale, Raphaël Blanchard, Archives de Parasitologie, Paris, scientific internationalism.
In 1930, the distinguished American entomologist
Leland Ossian Howard drew up a list of the most
important entomological events and discoveries of
the late nineteenth century. The first three dealt with
threats to agriculture – the gipsy moth, target of the
first federal quarantine against an insect on US soil;
the expansion of the San José scale through California’s fruit tree industry, which had elicited bans on
the importation of American plants from Canada
and Germany; and the cotton boll weevil, which had
apparently crossed the border from Mexico into
Texas before spreading eastward into Louisiana and
Mississippi. The fourth major event was the emergence of Medical Entomology.
For Howard, the study of mosquitoes and their
role in human diseases formed the epicenter of Medical Entomology. But how had this discipline begun?
Howard’s construction of Medical Entomology’s
genealogy was highly selective and inspired by the
success of medical parasitology, a vastly larger field.
Howard noted Patrick Manson’s demonstration of
the mosquito-borne nature of filariasis and cited
work on tick-borne Texas cattle fever. Above all, he
credited Ronald Ross’s discovery of the mosquitoborne nature of malaria as the foundation of the
emergent discipline.
Howard then imagined a future of steadfast and
seemingly inevitable cooperation between entomologists and physicians. In this context, he claimed Parasitology as a model for medical entomology’s bright
future and quoted the enthusiastic and inclusive
vision of a Parisian professor of Parasitology,
Raphaël Blanchard, who had written:
“The rapid movement which leads medicine into the
current of Parasitology cannot be stopped. In reality
these two branches of General Biology seem more or
Correspondence: Michael A. Osborne, Department of History,
University of California, Santa Barbara, CA, 93106-9410,
USA, Tel ++1 805 893 2901, Fax ++1 805 893 8795, e-mail:
osborne@history.uscb.edu
less distinct, but, as two rivers whose waters meet and
flow side by side for a certain distance soon come
together, so parasitology may include almost the
entire domain of medicine”1.
In comparison to the Nobel laureate Ronald Ross or
Patrick Manson, Blanchard is mentioned infrequently, if at all, in discussions of the new Tropical Medicine. Yet he institutionalized the new medicine,
developed teaching materials and texts and constructed a historical and progressive lineage for Parasitology. Blanchard had once hosted Howard in
Paris and authored many studies of Medical Entomology, including a massive Les moustiques: histoire naturelle et médicale (1905) and L’insecte et
l’infection; histoire naturelle et medicale des arthropodes pathogenes (1909)2. Both books and an earlier collaborative work with Alphonse Laveran on the
blood parasites of humans and animals reflected
taxonomic zeal 3.
Text books are generally not exciting pieces of scholarship. Yet as Ross ferreted out the lifecycle of malarial parasites in India he did so with the aid of Blanchard’s book on Medical Zoology 4. Blanchard cham1
L[eland] O[ssian] Howard, “Striking entomological events
of the last decade of the nineteenth century”, The Scientific
Monthly 31, no. 1 (July 1930): 5-18, quote on p 18. Provenance and translation unverified.
2
LO Howard, Fighting the insects; the story of an entomologist, telling of the life and experiences of the writer (NY:
The MacMillian Company, 1933), p 242; Raphaël Blanchard,
Les moustiques: histoire naturelle et médicale (Paris: FR de
Rudeval, 1905), idem, L’insecte et l’infection; histoire
naturelle et medicale des arthropodes pathogenes (Paris:
Librairie scientifique et litteraire, 1909).
3 A[lphonse] Laveran and R Blanchard, Les hématozoaires
de l’homme et des animaux, 2 vols (Paris: Rueff et Cie,
1895).
4 W[illiam] F Bynum and Caroline Overy, eds. The Beast in
the Mosquito: The Correspondence of Ronald Ross and
Patrick Manson (Amsterdam and Atlanta: Rodopi, 1998), pp
30, 52, 58, 79, 128, 312, 358, 436. The book was R Blanchard, Traité de zoologie médicale, 2 vols (Paris: J-B Ballière,
1889-1890).
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M.A. Osborne - Blanchard and the positioning of Medical Entomology in Paris
pioned Medical Natural History and Parasitology,
rather than Medical Entomology per se or the related
activities of “Medical Ecology,” a term sometimes
applied to Charles Nicolle’s 1909 discovery of the
relationship between the louse and epidemic typhus,
or “economic entomology,” a term sometimes favored
in Great Britain 5. Parasitology was Blanchard’s special
passion, and it mattered little to him whether leeches
or insects hosted and delivered the pathogenic organisms. By the end of his career he had authored more
than five hundred articles and books, and about fifty
of those were on leeches and some thirty on Diptera 6.
The consummate Parisian academic, Blanchard
was erudite, multi-lingual, cosmopolitan, and mainly a man of the classroom, taxonomy lab, and
writer’s study. Although he advocated a marriage of
laboratory and field studies and traveled to Algeria
on a natural historical mission in the spring of 1888,
he was not cut from the same cloth as many central
players in British tropical medicine such as Ronald
Ross, who had investigated tropical diseases on site
in India and West Africa, or Patrick Manson, who
had labored in Taiwan and China. When Blanchard
traveled abroad, he most often examined scientific
and medical institutions, or, once established, participated in scientific congresses around the world.
Among other things these travels resulted in a book
on medical inscriptions on historical monuments.
Around 1900, when Blanchard’s interests turned
more fully toward the European colonies and their
diseases, his student, Émile Brumpt, and not he,
traveled to Africa to investigate the relationship
between sleeping sickness and the tse-tse fly 7. The
dangers of the field avoided by Blanchard were very
real, and Brumpt, who crossed Africa from the Red
Sea to the Atlantic shore as the physician, naturalist, and photographer of the Bourg de Bozas expedition from 1901 to 1903, remarked on being bitten
by venomous ticks, saw many of the expedition’s
porters endure malaria, and recorded how many of
the group’s camels died after being attacked by tsetse flies 8. How then did Blanchard contribute to
medical entomology, parasitology, and the newlyemergent tropical medicine?
The historian of medicine Jean Théodoridès once
characterized Blanchard as the “grand father” of
modern French parasitology, and he certainly merits
this accolade as the mentor to Brumpt. More recently, Annick Opinel has identified Blanchard’s activities at the Paris Faculty of Medicine as one of three
institutional nodes of French medical entomology,
the other two being the Institut Pasteur and the military and colonial medical services. By 1907 the latter would be centered in Bordeaux, a naval post
graduate school in Toulon, and at the army’s new
post graduate school in Marseilles, the École
d’application du service de santé des troupes colonials (Le Pharo) 9.
Blanchard was fundamental to French medical
entomology for at least three reasons. First, insectborne diseases such as malaria, sleeping sickness,
and yellow fever are notoriously disrespectful of
boundaries and require international efforts at control. Blanchard had much international scientific
experience in Germany, England, and elsewhere and
was a cautious advocate of scientific internationalism and international standards for taxonomy at a
time of escalating tensions between Germany and
France. Second, Blanchard organized the teaching
and funding of parasitology and medical entomology
in the French capitol. He challenged the perceived
orthodoxy and traditions of the Paris Faculty of Medicine and was an able innovator in this largest and
most ossified of all French medical schools. Finally,
Blanchard was rooted in the broad traditions of a naturalist who synthesizes information and hazards
informed generalizations between diseases and organisms. The most visible French champion of parasitology, he was steeped in medical humanism and presented parasitology to the French learned community
in a non-threatening manner, positioning it as something that followed naturally from bacteriology and
represented the most recent stage in the natural progression of scientific medicine.
Raphaël Anatole Émile Blanchard was born in
1857 in Saint-Christophe (Indre-et-Loire) 10. Only
thirteen when hostilities opened in the FrancoPrussian War, he had a prodigious appetite for
5 Hervé Harant, “Cinquante ans de Parasitologie de langue
française”, Annales de Parasitologie 43, no. 1 (1968): 105115, p 110 for “medical ecology”. For the British context see
JFM Clark, “Bugs in the System: Insects, Agricultural Science,
and Professional Aspiration in Britain, 1890-1920”, Agricultural History 75, no. 1 (2001): 83-114.
6 G Lavier and J Théodoridès, “Raphaël Blanchard (18571919), médecin, naturaliste, et fondateur de la Société
Française d’Histoire de la Médecine”, Historie de la Médecine
7 (1957): 75-82, p 76.
7 On Brumpt see LW Hackett, “Émile Brumpt, 1877-1951”,
The Journal of Parasitology 38, no. 3 (June 1952): 271-273,
and Henri Gaillard, “Émile Brumpt”, Dictionary of Scientific
Biography, vol 2, pp 533-534.
8 Émile Brumpt, Mission de Bourg de Bozas de la Mer
Rouge à l’Atlantique à travers l’Afrique tropical, conference
faite à la Société de Géographie le 5 juin 1903 (Paris: FR de
Rudeval, 1903).
9
Jean Théodoridès, “La contribution française à la parasitologie médicale et à la pathologie exotique de 1900 à
1950”, Histoire des Sciences médicales 27, no. 3 (1993): 223231, quote on p 224. Annick Opinel, “The emergence of
French medical entomology: the respective influence of universities, Institut Pasteur and army physicians (1890 to ca
1938)”, draft typescript generously shared by Dr Opinel.
10
For biographical information see André Cornet, “Raphaël
Blanchard”, http://www.bium.univ-paris5.fr/sfhm/histoire2.htm
[accessed 10 Jaunary 2008], “Raphaël Blanchard (18571919)”, http//:www.pasteur.fr/infosci/archives/blr0.html
[accessed 10 January 2008], Émile Brumpt, “Raphaël Blanchard”, Archives de Parasitologie 16 (1913-1919): i-iv, and G
Lavier and J Théodoridès, “Raphaël Blanchard (1857-1919)”.
Also of use is the Blanchard collection at the Institut Pasteur,
FR IP BLR. I wish to thank Stéphane Kraxner of the Institut
Pasteur’s Service des Archives for his good humor and for
guiding me through this and other collections.
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215
learning and accumulated impeccable scientific credentials. Few Frenchmen of his generation combined his skills at histology, developmental biology,
microscopy, and medical administration. Upon
arrival in Paris in 1874, he sampled courses at the
Faculty of Medicine and was struck by those of
Charles Robin. His early interests, extremely varied,
were in histology and developmental biology rather
than in entomology. While working in Robin’s laboratory of zoological histology at the Sorbonne’s
École pratique des Hautes Études from 1875 to
1877, he became friends with the laboratory’s assistant director, Georges Pouchet, and continued medical studies. Blanchard became Pouchet’s protégé
and learned something of experimental teratology
after Pouchet began teaching zoology at the École
Normale Supérieure. He soon won a fellowship and
spent a year in Germany and Austria. It was the first
such fellowship after the Franco-Prussian War. If
Blanchard had doubts about going to Germany, his
mother had even more until Pouchet convinced her
it was the best thing for young Raphaël’s career 11.
Blanchard studied embryology in Vienna at
Samuel Leopold Schenk’s Institute of Embryology.
He also traveled to centers of the new physiology,
notably to Leipzig where he visited the new anatomical institute of Wilhelm His. In 1880, the same year
he defended his medical thesis on nitrous oxide and
the anesthetic methods of Paul Bert (in whose laboratory he then worked) he returned to Germany and
this time also traveled to Russia and Scandinavia,
publishing excerpts on his travels in Progrès médical 12. Blanchard was impressed with the material
resources lavished on German science and returned
to Paris admiring some German scientists, and disliking others. Other French scientists and physicians
had arrived at similar conclusions but Blanchard
went further than most by authoring a glossary of
German and French anatomical and zoological
terms 13. He would be suspicious of Germany and
German science throughout his life, and later
clashed with German taxonomists over the use of
Latin in classification. But cognizance of German
scientific methodology was only one of many influences informing Blanchard’s work on Medical Entomology. He was thoroughly French and especially
prominent in Parisian scientific societies.
As a young man he had been a savior of sorts for
the Société zoologique de France, a group founded
in 1876 by Aimé Bouvier, a merchant of natural historical materials, which included a coterie of amateur naturalists, and a sprinkling of aide-naturalistes
from the Muséum. Bouvier soon resigned in disgrace and the group’s treasury was short the considerable sum of 5,000 francs 14. Blanchard brought
stability to the group as its general secretary from
1879 to 1900, and eventually served as its president. Even before he gained a professorial post at
the Faculty of Medicine, Blanchard had accumulated much international experience, taxonomic expertise, and talents at negotiation and organization. He
would need and use all of these in his work on Parasitology and Medical Entomology.
11
R Blanchard, “Souvenirs d’Allemagne”, Bulletin de la
Société zoologique de France 40 (1915): 3-26, p 8.
12
R Blanchard, Les universités allemandes (Paris: A Delahaye & E Lecrosnier, 1883).
13
R Blanchard, Glossaire allemande-français des terms
d’anatomie et de zoologie (Paris: Asselin et Houzeau, 1908);
Harry W Paul, The Sorcerer’s Apprentice: The French Scientist’s Image of German Science, 1840-1919 (Gainesville: University of Florida Press, 1972).
14 Robert Fox, “La Société zoologique de France. Ses origines
et ses premières années”, Bulletin de la Société zoologique de
France 101[5] (1976): 799-813.
15 R Blanchard (ed), Congrès International de Zoologie, Paris
1889. Compte-rendu des séances (Paris: Société zoologique de
France, 1889).
16 Ibidem, pp 313-314.
17 R Blanchard, “Souvenirs d’Allemagne”, pp 19-21, quoted
in R Fox, op cit, p 807.
Blanchard, taxonomy, and internationalism
Blanchard’s international reputation and his tense
relationship with German taxonomists were evident
in 1889 when Paris hosted the first International
Congress of Zoology in conjunction with that year’s
Exposition Universelle. Blanchard was friends with
Alphonse Milne-Edwards, the Muséum national
d’histoire naturelle’s Professor of Mammals and
Birds who became the President of the Congress.
The established Muséum Professors, Édmond Perrier and Léon Vaillant, filled the vice-presidentships.
Most of the organizational work of the congress and
the editing of the proceedings, however, fell to Blanchard who cited himself more than any other participant in the proceedings’s index. Blanchard was
especially prominent in the two largest sections of
the congress, the section on the geographical distribution of animals, and the section on zoological
nomenclature 15.
Zoology lagged behind Geology and Botany in
developing an agreed-upon nomenclature. Blanchard lobbied congress participants to adopt the
binominal nomenclature and to use Latin rather
than a national language for all matters, even for the
names of scientists if they figured in the names of
new organisms. But no agreement was reached at
Paris or the subsequent congress in Moscow. At
Paris Blanchard had intervened in disagreement
with the Berlin naturalist Schiller-Tietz’s paper on
the general laws of Parasitology 16, and at the third
international zoological conference held at Leyden
in 1895, he accepted the presidency of the Permanent International Commission on Zoological
Nomenclature. He repeatedly disagreed with German scientists over nomenclature and the use of
national languages, particularly after the Berlin zoological congress of 1901, and he finished his career
by lashing out at the German nation and its “barbarian hordes”17. The fruit of his efforts, however,
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was an agreement on zoological nomenclature
adopted at the Bern conference of 1904 and published the next year.
In the same year Blanchard published Les moustiques: histoire naturelle et médicale, his major
study of medical entomology. The work reflected
both his international connections and his taxonomic zeal. In the preface he thanked Howard for his
assistance with numerous illustrations. He also
thanked Frederick Vincent Theobald, Chief of the
British Museum’s Entomology Section. In Great
Britain Colonial Secretary Joseph Chamberlain’s policy of “constructive imperialism”, backed by funding
from the Colonial Office, was energizing a whole
range of tropical sciences from medicine to entomology, Theobald, a renowned expert on entomology and mosquitoes, was compiling a multi-volume
study of Culicidae 18. Theobald sent Blanchard several specimens classified by himself, and Les moustiques drew mainly on them and on a collection
assembled by Blanchard in his Paris laboratory.
Blanchard’s book was less comprehensive than
Theobald’s, and he addressed mainly physicians
serving in regions of the world afflicted by malaria
and yellow fever. His book skillfully interwove chapters on taxonomy and mosquito anatomy with chapters on prophylaxis, epidemiology, and parasitology.
In terms of taxonomy, Les moustiques assiduously
followed the new International Rules of Zoological
Nomenclature of which Blanchard had been a major
architect 19.
Thus by the first decade of the twentieth century,
Blanchard was fully engaged in medical entomology,
but not to the exclusion of other activities. Through
pamphlets and other venues, he implored travelers
and military men bound for the tropics to capture
mosquitoes, biting flies, ticks, and other organisms
and instructed them on how to pack them in matchboxes or other containers and send them to his laboratory 20. At decade’s end he traveled to Brussels for
the first International Congress of Entomology to
deliver an address entitled “Entomology and Medicine”, a contribution ordered by various taxonomic
groups dealing mainly with arthropods. The talk
traced the etiology of several insect and bug borne diseases and spoke against miasmatic theory while speculating on the causes of beriberi and other human and
animal afflictions. If the causes of European diseases
JFM Clark, “Bugs in the System”, (note 4), p 84. Frederick Vincent Theobald, A monograph of the Culicidae, or mosquitoes. Mainly compiled from the collections received at the
British museum from various parts of the world in connection
with the investigation into the cause of malaria conducted by
the Colonial office and the Royal society, 5 vols, atlas (London: Printed by order of the Trustees, 1901-10).
19 R Blanchard, Les moustiques, “Préface”, pp v-vi, and
Règles internationales de la nomenclature zoologique adoptées par les congrès internationaux de zoologie. International
rules of zoological nomenclature. Internationale regeln der
zoologischen nomenklatur (Paris: FR de Rudeval, 1905).
20 R Blanchard, Instructions sommaires pour les pays
chauds (Paris: FR De Rudeval, 1905).
18
remained unknown, he speculated on their etiology
and drew analogies with tropical afflictions which
seem to produce similar symptoms. Parasitology stood
clearly at the pinnacle of the medical art. Formerly it
had been thought “that bacteriology was going to be
the last word of medicine; it is now outstripped and
considerably so by animal parasitology and medical
entomology, the sphere[s] of which are truly without
limits”. We can predict, he continued, a new golden
age for humanity where henceforth “immense territories open up to the white race where formerly, until
now, it was stalked by a hundred unknown enemies
now revealed. These discoveries, and those of tomorrow, will change the face of the world”21.
Parasitology at the Faculty of Medicine and
the Institute of Colonial Medicine
In 1883, Blanchard passed the aggregation in natural
history at the Paris Faculty of Medicine and began
teaching medical zoology where he seconded the
botanist Henri Ernst Baillon who had held the Chair
of Medical Natural History since 1863. Baillon’s
interests lay in general and taxonomic botany, and
secondarily in materia medica. Blanchard focused resolutely on medical zoology, but so had Baillon’s predecessor, Charles Alfred Moquin-Tandon. Blanchard’s
interests and erudition resembled those of MoquinTandon in several ways, and both men provided medical students with a respectable knowledge of a larger scientific culture and tradition. The naturalist and
prolific entomologist and science writer, Jean-Henri
Fabre, described Moquin-Tandon as “a naturalist
with far-reaching ideas, a philosopher who soared
above petty details to comprehensive views of life, a
writer, a poet who knew how to clothe the naked
truth with the magic mantle of the glowing word” 22.
Blanchard and Moquin-Tandon also shared a love
of the arts. Blanchard was attracted by the painting
of his era, photography, history, vernacular architecture and sundials 23. Moquin-Tandon inclined toward
poetry which he composed and published in his
native provençale language. Additionally, both men
published on leeches and were fascinated by these
curious animals 24. Blanchard wrote at a time of high
imperialism authoring studies of both French and
North African leeches 25. Finally, Moquin-Tandon,
21
R Blanchard [résumé by J Desneux], “L’Entomologie et la
Médecine”, 1 Congrès International d’Entomologie, Bruxelles,
1-6 aôut 1910 (Bruxelles: Hayez, 1912), vol 1, Historique et
process-verbaux, pp 113-123, quotations on pp 122-123.
22
Quote from The Life of the Fly (1913) reproduced in
Plant Talk, http://www.plant-talk.org/pages/15fabre.html
[accessed 10 January 2008].
23
R Blanchard, L’art populaire dans le Briançonnais (Paris:
É Champion, 1914).
24
Erwin H Ackerknecht, Medicine at the Paris Hospital
(Baltimore: The Johns Hopkins University Press, 1967), ch 6,
“Broussais”, pp 61-80.
25
Henri Gadeau de Kerville, Voyage zoologique en
Khroumirie (Tunisie) mai-juin 1906 (Paris: J-B Baillière, 1908).
Blanchard examined the leech specimens from the voyage.
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like Blanchard after him, signaled the utility of medical natural history for the colonial enterprise and
identified the horse leach as one of the main causes
of disease in Algeria 26. It was Blanchard, however,
who created enduring institutions which made Medical Entomology and Parasitology relevant to colonial development.
Blanchard strove to make Natural History relevant
to Medicine. Early lectures at the Faculty of Medicine focused on the life cycles of parasites. It was,
as he never tired of claiming, the first such comprehensive class ever given at a French medical faculty.
When Baillon fell dead in the Faculty’s botanical
garden in July of 1895, it was logical for Blanchard
to assume the Chair. But the Chair itself, so long
associated with descriptive botany, now seemed
redundant and marginal to medicine. The reasons
for this were various but due mainly to reforms in
medical and university studies in the early 1890s,
especially the reform of 1893. New students bound
for medical careers were required to complete a
preparatory year in the physical, chemical and natural sciences. Descriptive botany was now a premedical subject to be taught at faculties of science
rather than in medical schools.
Other problems internal to the Faculty of Medicine also frustrated Blanchard. Prior to the reforms,
Medical Natural History had been taught in the first
year of medical studies. But this was the age of Pasteur and Koch, and with Baillon’s death the Faculty
stopped maintaining its botanical garden and Baillon’s classes went untaught for two years. No longer
was Natural History taught as a unified whole, as a
foundation of Medicine and perhaps as a reason for
etiology’s claims to scientific status. Reforms now
split the subject between third and fourth year
courses. Finally, in 1897, the Paris Faculty of Sciences claimed the medical faculty’s botanical garden. Clearly, new directions and visions were
required. For Blanchard, that vision was parasitology.
In articles and a multitude of brochures, Blanchard railed against those who termed Parasitology
an accessory to medical practice. He portrayed the
new discipline as mounting a “…a frontal attack on
the most recalcitrant questions of hygiene and
pathology[;] it brings to diagnosis the precision
which it too often lacks, casts light on morbid etiology and prophylaxis, and explains symptomatology
and pathology” 27. One issue, of course, was that the
global distribution of pathogenic human parasites
was skewed toward the tropics, not Europe, and
Paris was not a center for the study of these sorts of
diseases and organisms. In fact, Blanchard had been
interested in the French colonies for years and orga26 Alfred Moquin-Tandon, Elements of Medical Zoology
(London: H Baillière, 1861), ch 4, “Leeches”, pp 137-147, p
217.
27 R Blanchard, “Réorganisation des études médicales; le
PCN”, pp 485-486, p 486.
217
nized many popular lectures on the Natural History
and Anthropology of North Africa and Madagascar 28. But there was competition from several quarters, and as noted earlier by Opinel, training in colonial medicine could be had at a number of locations
including the Institut Pasteur, or at institutions run
by the navy and army and centered in cities which,
unlike Paris, were heavily invested in colonial activities: Bordeaux, Toulon and Marseilles. Blanchard
finally gained the Chair of Medical Natural History
on 25 July 1897 and by November of 1906 had convinced his colleagues to rename his post the Chair
of Parasitology and Medical Natural History. This
change of title in Paris, according to Blanchard, ended definitively the teaching of purely descriptive
Zoology and Botany at the Faculty. Yet in terms of
the formal recognition of parasitology, Paris was a
few years behind the faculty of medicine at Lille
where in 1894 Alfred Giard had become France’s
first incumbent of a Chair of Parasitology 29.
Professional and patriotic urges gave rise in 1902
to the Faculty’s new Institute of Colonial Medicine.
This institution, and Blanchard’s own laboratory,
would prove beyond a doubt the utility of his specialism for medical education. Still, the Faculty of
Medicine was more reserved on Parasitology, and
waited until 1909 to oblige its third year students to
take a laboratory-based course on the subject and an
examination 30. Most certainly, the Institute of Colonial Medicine gained Blanchard an audience of students, particularly international students.
Blanchard modeled his Institute of Colonial Medicine, which he directed until his death in 1919, on
the tropical medical schools of Manson and Ross at
London and Liverpool, respectively. The Paris Institute was also a post-doctoral school where about
thirty physicians followed three months of classes to
obtain the diploma of colonial physician. While the
London School had three separate three-month long
sessions in 1899-1900, the Paris Institute would
have but one three-month session per year 31. Blanchard arranged for a clinic for tropical diseases at a
hospital run by the Association of French Women in
Auteuil. He also gained promises for 150,000 francs
of funding from the governor of French Indo-China,
who subsequently reneged, and agencies who did
not; the Union coloniale française, the Minister of
the Colonies, and the government of Madagascar.
28
See for example the collected work Madagascar au début
du XX siècle (Paris: FR de Rudeval et cie, 1902).
29
Georges Barrière, “Raphaël Blanchard (1857-1919), sa
vie, son oeuvre” (Thèse pour docteur en médecine, Université
de Aix-Marseille [II], 1982), p 28. Cf G Lavier and J Théodoridès, “Raphaël Blanchard (1857-1919)”, p 77, note 2.
30
R Blanchard, “Projet de réorganization du Service de la
Parasitologie”, Archives de Parasitologie 13 (1908-1909):
311-342, p 311.
31
R Blanchard, “L’enseignement de la médecine tropicale”,
Le Progrès Médical (15 juillet 1899): 38-42; idem, “La
médecine des pays chauds. Son enseignement, ses applications
à la Colonisation”, Le Progrès Médical, 3e série, tome X, no.
44 (4 novembre 1899): 289-293.
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Blanchard’s twenty-one lectures and associated practical exercises on Parasitology anchored the curriculum. These were supported by clinical rounds and
lectures on Bacteriology, Exotic Pathology, Tropical
Hygiene and Climatology, and Dermatology. By
1910 the Institute had graduated 249 students in
nine classes; about half of these were French (110),
and a third (72) hailed from Latin America.
The Institute focused on the research and teaching
of “Exotic Pathology” and included Medical Entomology in course content 32. Blanchard had good technical skills, and he continually refreshed his knowledge at conferences and by attending the microscopy
course taught at the Institut Pasteur in 1896. But his
classes at the Institute and Faculty of Medicine followed his proclivities and they tended to be general
and even philosophical exercises. For example, his
Parasitology course included twenty-one lectures
and practical demonstrations, and the very first lesson covered insect vectors as one of four modes of
transmission. Subsequent lectures treated malaria,
Texas cattle fever, trypanosomes and sleeping sickness, and filarial afflictions. He also devoted a lecture to parasitic insects, especially those encountered in the colonies.
By 1908 what we now identify as medical entomology was in fuller evidence at the Institute in the
laboratory course of Maurice Langeron, a mycologist who undertook medical study in Dijon and
Paris. Like Blanchard, Langeron also traveled to
Germany to perfect his German language skills.
Upon returning to Paris he became Blanchard’s secretary at a salary of fifty francs per month and
would eventually head Blanchard’s laboratory and
work tirelessly on Blanchard’s journal, the Archives
de Parasitologie. Langeron’s laboratory course of
1908 covered microscopy and staining techniques,
the diagnosis of blood disorders and bacteriological
and mycological diseases. About a dozen laboratory
exercises and preparations dealt with insects or diseases carried by insects, especially malaria, but also
sleeping sickness, tick fever, and elephantiasis. Students were also instructed on how to identify the
female Culex mosquito and its larvae, the Anopheles mosquito, and had prepared slides on Plasmodium malariae, Plasmodium vivax, and three slides
on Plasmodium falciparum 33. In sum, Langeron’s
course was, like Blanchard’s, an overview of the
interactions of parasites and humans with medical
entomological topics placed there in.
Most certainly, numerous students and researchers
who later became professors of Parasitology or Tropical Medicine at provincial and foreign medical fac32 R Blanchard, “L’Institut de Médecine Coloniale, histoire
de sa foundation”, Archives de Parasitologie 4 (1902): 585603, p 585.
33 Maurice Langeron, “Technique des manipulations complémentaires de parasitologie”, Archives de Parasitologie 12
(1908): 177-191. On Langeron see Johanna Westerdijk and
Jacomina Lodder, “Maurice Langeron, 1874-1950”, Antonie
van Leeuwenhoek 17, no. 1 (December 1951): 275-277.
ulties trained at the Institute. This included Blanchard’s first student, Jules Guiart, who in 1906 would
become Professor of Medical Natural History (Professor of Parasitology after 1907) at the Lyon Faculty
of Medicine. In 1912 Blanchard brought a young
physician from Nancy to Paris, Charles Joyeux, who
arrived with substantial experience in West Africa and
Upper Guinea. After service on the front in World
War I he returned to Paris and passed the aggregation
in 1920. In 1930 Joyeux became the first professor of
Parasitology at the new Faculty of Medicine in Marseille 34. In addition to institutionalizing academic parasitology in Paris and founding the Institute of Colonial Medicine and the training many students, Blanchard promoted and presented Parasitology to a
wider public through his historical activities and
through the pages of the Archives de Parasitologie.
Parasitology, Medical Humanism, and History
The entrepreneurial Blanchard promoted Medical
Parasitology and thus medical entomological study
through several venues. I should like to conclude
with his broader historical and cultural activities
and note how these intersected with Parasitology
and positioned his relatively new science in the lineage of human knowledge. Blanchard routinely
included historical articles, photographs of monuments and people, artistic caricatures, and similar
items in the Archives de Parasitologie, a journal he
founded in 1898 and edited until the outbreak of
World War I. Additionally, historical examples were
plentiful in his teaching. For example, his course on
Parasitology at the Faculty of Medicine in 19101911 covered the sociology of parasitism and its
role in ancient Greece and Rome, while a course on
parasitism and infection the next year examined
ancient theories of Parasitology, and covered what
he termed the precursors of modern scientific parasitology from the seventeenth century Italian physician, Francesco Redi, to more modern figures
including Vincent Raspail and Louis Pasteur 35. All
of these researchers seemingly contributed to the
logical progression of Parasitology which now
crowned the medical sciences.
At the turn of the last century, as sectors of the
healing arts became more scientific and medicine
more specialized, the perceived canon of medicine
grew so vast as to be beyond a single person’s mastery. A generation of men such as the Canadian
physician William Osler, and Blanchard, sought presentation of themselves as learned men rather than
narrow technicians or engineers. Osler collected
medical and scientific books, wrote medical history,
and delighted in medico-literary pranks. Blanchard
dearly wanted to be a man of “haute culture”, and
34 G Barrière, “Raphaël Blanchard (1857-1919), sa vie, son
oeuvre”, pp 28-30.
35 “M le Professeur R Blanchard. Cours de Parasitologie”,
Archives de Parasitologie 15 (1911-1912): 328-330.
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from his facility in foreign languages, to his photographic activities and commentaries on art history
and collecting, he was exactly that. In the sixteenth
and last volume of the Archives de Parasitologie,
which appeared in 1919, Blanchard reflected on the
historical, cultural and artistic dimensions of Parasitology in a section of the journal entitled “Parasites
and parasitological illnesses in history, poetry, and
art” which reproduced comments by Thucydides on
the plague of Athens and eleven photographs of
paintings of Napoleon in Egypt. Since the foundation of the journal in 1898, he wrote, it had been his
intention to soften the severe and narrow character
of which scientific journals ordinarily suffered, and
to intercalate historical and artistic content with scientific articles, rather than grouping non-scientific
pieces in a single volume 36.
In reviewing the various historical and artistic
contributions appearing in the pages of the Archives
de Parasitologie, one is struck by the number of
documents and photographs of items relating to
Louis Pasteur, who had died in 1895. Included
throughout the volumes of the early 1900s are photographs and an account of the dedication of a statue erected in Paris in 1904, a photograph of a monument to Pasteur in an eponymous village in Algeria, and other photographs of the well-known statues at Melun, and at Arbois, where tablets around
the base of the statue show Roux and Pasteur providing anti-rabies vaccine and the assistance Pasteur’s science provided to agrarian France. One
image from 1910, that of a painted earthenware
plate with scenes of Pasteur in his laboratory and
watching over the anti-rabies vaccination of a young
boy, strikes modern readers as particularly strange.
Yet these cultish remembrances of Pasteur served
Blanchard’s purposes by connecting Parasitology
with popular culture and utilitarian traditions and
positioned Blanchard’s science as the next logical
step in a narrative of the progress of science. But
they were also quite in line with Blanchard’s views
on what constituted parasitological information, and
how history informed his science.
In addition to his historical work published in the
Archives de Parasitologie, Blanchard is remembered
as the founder of the Société française d’histoire de
la médecine in 1902. One historical project was similar to his catalog of sundials and collected a total of
1,258 inscriptions on monuments of importance to
Medical History. Blanchard, with the encouragement
of Karl Sudhoff, a historian of medicine at the University of Leipzig, focused on early modern and
modern monuments prior to 1900. He collected
much of the information during professional conferences and while traveling, and his taxonomic hand
36
R Blanchard, “Les parasites et les maladies parasitaires
dans l’histoire, la poésie et l’art”, Archives de Parasitologie 16
(1913-1919): 579-580, which introduces a lengthy section on
plague in Athens and Marseille, and in the writings of Thucydides and La Fontaine, pp 581-637.
219
was at work in much of the translation of the entries
arranged by the name of the person commemorated
or the structures upon which the text appeared 37.
It is easy to charge that Blanchard was antiquarian in his methods. But his historical work, so often
directed at the pre-history and history of Parasitology, spoke to an important issue of scientific
methodology, that of inclusiveness after the fashion
of ethnography or the naturalist who synthesizes
vast amounts of information from diverse sources 38.
Evidence of disease, and its past ravages and
appearances, could be found in numerous sources,
and Blanchard wanted his colleagues to account for
them and to consider artistic, photographic, literary,
and cultural depictions as evidence and possibly to
bring these to bear on diagnosis. Even Émile
Brumpt, whose knowledge joined most effectively
the discoveries of the laboratory and field, would
use linguistic evidence in estimating the prevalence
of sleeping sickness, concluding that if the residents
of the Congo basin had no special name for the disease, it was likely rare or of recent occurrence 39.
Conclusion
The Archives de Parasitologie suspended publication
in 1914 and when Blanchard died on 7 February
1919 the Archives, after a final issue carrying his
obituary, perished with him. A complete list of war
casualties might well include the inclusive internationalism and medical humanism typified in Blanchard’s science. In comparing the Archives de Parasitologie with the first few issues of the Annales de
Parasitologie Humaine et Comparée, begun in 1923
by Émile Brumpt and edited by him, it is apparent
that the newer publication had no interest in historical articles or poetry. Whereas Blanchard had welcomed contributions in French, German, English,
Spanish or Italian, and had dedicated the Archives
to the “study of parasites, envisaged in their most
diverse aspects”, the newer journal accepted only
French language contributions and narrowed its
brief to original research and taxonomy. Medical
entomology had now come of age and had a clear
utility, and while Brumpt did not immediately use
this terminology, he did mention insects and malaria in his inaugural editorial 40.
Other facets of Blanchard’s program survived the
war. His Institute of Colonial Medicine pointed
37 Cf André Cornet, “Raphaël Blanchard”; R Blanchard,
Épigraphie Médicale: Corpus inscriptionum ad medicinam
biologiamque spectantium (Paris: Asselin et Houzeau, 1909).
38 Paul L Farber, Finding Order in Nature: The Naturalist
Tradition from Linnaeus to EO Wilson (Baltimore and London: The Johns Hopkins University Press, 2000).
39 Émile Brumpt, “Maladie du sommeil, distribution géographique, étiologie, prophylaxie”, Archives de Parasitologie 9
(1905): 205-225.
40 Compare, for example, R Blanchard, “Notre programme”.
Archives de Parasitologie 1 (1898): 5-7, quote on p 6, with É
Brumpt, “Avant-propos”, Annales de Parasitologie Humaine
et Comparée 1 (1923): 1-3.
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toward the new regime in the French colonies,
where civilian healers replaced those formerly
trained by the navy. This civilianization of colonial
medicine did not work out quite as planned. Not
many directors of the Pasteur Institutes bothered to
complete the Institute diploma and even in the new
colony of Viet-Nam, those who trained first at Bordeaux, and took post-graduate training either at the
naval school in Toulon, or the Pharo army school in
Marseille, had inside tracks on colonial careers. Yet
in the larger scheme of things, Blanchard’s colonial
turn was prescient as was his cluster of medical concerns, which included anthropology and race as factors in disease and illness. Knowledge of Parasitol-
ogy, Medical Entomology, colonial or exotic diseases, and the health of colonized peoples in
Europe, was valued during the war and had been
useful to medical management of the half million
troupes indigènes, the force noir, who came to the
assistance of France in its hour of need 41.
41 These issues are examined in greater detail in Michael A
Osborne and Richard S Fogarty, “Views from the Periphery:
Discourses of Race and Place in French Military Medicine”.
History and Philosophy of the Life Sciences 25 (2003): 363389. See also Richard S Fogarty, Race & War in France: colonial subjects in the French Army, 1914-1918 (Baltimore: The
Johns Hopkins University Press, 2008).
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The Sergent brothers and the antimalarial campaigns
in Algeria (1902-1948)
J.-P. Dedet
Université Montpellier 1, CHU de Montpellier, Laboratoire de Parasitologie, Montpellier, France.
Abstract. Edmond and Etienne Sergent, “the Sergent brothers”, were both born in Algeria. They both
studied medicine at the Algiers Medical School and then followed the Course of Microbiology of Emile
Roux at the Institut Pasteur in Paris (1899-1900). From 1900, they were put in charge of a permanent
mission aimed at antimalarial control in Algeria, which was supervised by the Institut Pasteur. The first
campaign was carried out during the summer of 1902 at a station of the East Algerian Railway Company. The success of this mission lead to the creation of the Antimalaric Department of Algeria in 1904,
which was directed by Etienne Sergent for the duration his life. This antimalarial programme was progressively extended to many other locations. The programme was optimized between 1927 and 1947,
in the experimental field study of the Ouled Mendil Marsh, where global environmental measures and
drainage lead to settlement of farms, the families of which did not suffered from malaria. At a time when
neither insecticides nor synthetic antimalarial drug existed, antimalarial control measures that were
developed tended to target human reservoirs and the mosquito vectors. The extension of the programme
across the Algerian territory lead to a decrease of both malaria endemicity and extension of affected
areas.
Key words: malaria, antimalarial campaigns, Sergent Edmond, Sergent Etienne, Algeria.
The work of Louis Pasteur generated not only the
creation of the Institut Pasteur, in Paris (1888), but
also a mundial dissemination of antirabic vaccination centres and microbiology laboratories, which
were gradually created throughout the French colonial empire, in French Indochina (Saigon, 1891, and
Nhatrang, 1895), in North Africa (Tunis, 1893,
Algiers and Tanger, 1910), in South Saharan Africa
(Dakar, 1896; Tananarive, 1898; Brazzaville,1908),
and also in independant countries such as in Constantinople (1893), Bruxelles (1900), Chengdu
(1911), Bangkok (1913), Athens (1920) and Tehran
(1920) (Dedet, 2000). During the 19th century,
four Pasteur Institutes were created in North Africa,
three of which are still active, including the one of
Algiers in which the Sergent brothers worked for all
of their careers.
The Sergent brothers, Edmond and Etienne, were
both born in Algeria, in Philippeville in 1876, and
in Mila in 1878 respectively. They both studied
Medicine at the Medical School of Algiers, and completed their training in Microbiology by following
the “Cours de Microbie technique” of Doctor Emile
Roux, at the Institut Pasteur in Paris. Edmond was
also trained in entomology by Louis Eugène Bouvier, from the Muséum national d’histoire naturelle of
Paris.
Edmond has been director of the Institut Pasteur
d’Algérie from 1912 to 1963, and Etienne directed
the Anti-malaric Department of Algeria from 1904
to 1948. During all these years of scientific activity,
they both devoted a large part of their
ical and Veterinary Entomology.
This paper focuses on the extensive
gent brothers carried out in malaria
and control, which have contributed
developments in this field.
work to Med-
In 1900, Edmond and Etienne Sergent started their
scientific career, just twenty years after the discovery of the malaria parasite, by Alphonse Laveran, in
Algeria (Laveran, 1880), and only two years after
Ronald Ross had demonstrated the transmission of
the sparrow malaria, Plasmodium relictum, by the
grey mosquito, Culex fatigans (reported by Manson,
1898-99). Ross’s discovery had been rapidly completed and extended to human malaria by Battista
Grassi and his colleagues, who showed the entire
sporogonic cycle in Anopheles claviger (presently
known as Anopheles maculipennis), in Italy (Grassi
et al., 1899). These discoveries opened the era of
antimalarial control, the first campaign being carried out during the summer of 1899 by Angelo Celli in Latium, Italy. Rapidly antimalarial campaigns
were generalized to Italy, and extended to the British
colonial empire. They were also the starting point of
all the work of Edmond and Etienne Sergent on
malaria epidemiology and control. In the French
colonies, the Sergent brothers were the first to
develop malarial control programmes.
Correspondence: Jean-Pierre Dedet, Laboratoire de parasitologie, 163 rue Auguste Broussonet, 34090 Montpellier, France, e-mail: parasito@univ-montp1.fr
The Sergent brothers showed that malaria in Algeria was due to three Plasmodium species: P. falciparum, P. vivax and P. malariae. Etienne Sergent
made a first observation on Anophelines by discov-
works of Serepidemiology
to significant
The Sergent brothers malaria approach
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J.-P. Dedet - The Sergent brothers and the antimalarial campaigns in Algeria
ering the presence of Anopheles larvae in Algiers in
October 1900 (Sergent Et., 1901b). He found two
species: Anopheles maculipennis and a new species
he sent to Theobald, who created for it the name of
A. algeriensis (Theobald, 1903). During the same
summer, Etienne Sergent observed the presence of
Anopheles larvae in the vicinity of Paris, an area
where malaria had disappeared a long time ago
(Sergent Et., 1901a). This observation lead the Sergent brothers to the elaboration of the concept of
“anophelism without malaria” (Sergent Edm. & Et.,
1903a).
From 1900, Edmond and Etienne Sergent were put
in charge of a permanent mission for antimalarial
control in Algeria, which was headed by the Institut
Pasteur of Paris. During the 10 years that followed,
Etienne was permanently working in Algeria, while
Edmond spent only summers and autumns there,
which were the malaria transmission periods. In a
few years, they demonstrated the presence of
Anophelines in all the localities of Algeria where
malaria was prevalent (Sergent Edm. & Et., 1903b).
They carried out their first experience of antimalarial methods during the summer of 1902, in a single
station of the East Algerian Railways Company: the
station of “L’Alma”, chosen for its high incidence of
malaria. The measures were only directed against
the mosquitoes: weeding and cleaning the larval
resting sites, spreading oil on them, closing doors,
windows and chimneys of the station with copperwire gauze (Sergent Edm. & Et., 1903c). These simple measures were so successful that the director of
the East Algerian Railways Company asked the Sergent brothers to extend the measures to seven of the
most infected stations. The 1903 campaign lead to
a decrease of malarial incidence in these railways
stations from 35% to only 6%.
The Antimalaric Department of Algeria
In 1904, the General Governor of Algeria, Auguste
Jonnart, created the Antimalaric Department of
Algeria, which the Institut Pasteur was asked to
direct. It was Etienne Sergent who was designated
to take this charge, a position which he kept all his
life. The aim of the Antimalaric Department of
Algeria included (i) experimentation, (ii) educational propagandism, and (iii) application of prophylactic measures.
The evaluation of the protective value of the available prophylactic measures, according to the different environments, was carried out in nine experimental field stations located throughout the three
algerian “Departements” of Oran, Alger and Constantine.
The success of the antimalarial measures in the
railways stations was given upon the basis of the
antimalarial education of rural populations. The
people daily passing in the stations could identify
the success in malaria prevention with the measures
taken in the stations, such as closing doors and win-
Figure 1. Poster on malaria for public information and education edited by the Institut Pasteur of Algeria. (Photo from
“Notice sur l’Institut Pasteur d’Algérie, Alger, 1949).
dows with copper-wire gauze. The Antimalaric
Department also issued post-cards, posters, and
small brochures for public information and education (Figure 1).
The application of the antimalarial measures was
preceded by an epidemiological study with mapping,
which determined the different endemic indices:
splenic, splenometric and plasmodic indices in
humans, sprozoitic index in sandflies, and during
which were detected the Anopheline breeding sites
(Figure 2).
The decision of initiating a campaign was depending on the request of the local communities, their
ability to provide a medical supervision, the results of
the epidemiological studies, and available funds.The
aim was to extend the antimalarial campaigns step by
step, gradually, from one area to another. The campaigns were restricted to the railways stations
between 1902 and 1904. They gradually, involved an
increasing number of villages since 1905. The numbers of population protected reached 32,000 by
1926. Within the nine experimental field stations, the
antimalarial measures were developed further and
their efficiency tested according to a variety of environmental factors (Sergent Edm. & Et., 1928).
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223
Figure 2. Map of malaria of the Mitidja plain, based on endemicity indices. (Photo from Sergent Edm. & Et., 1928).
Between 1927 and 1947, the Sergent brothers
developed a specific programme in an unhabited
area, the Ouled Mendil Marsh, where they developed an experimental field study based on global
environmental measures and drainage which lead to
settlement of farms, the families of which never suffered from malaria [Sergent Edm. & Et., 1947).
Entomological studies
During all these years of development of antimalaria campaigns, the Sergent brothers carried out regular observations on the morphology, biology and
ecology of the Anopheles species present in Algeria.
From a taxonomic point of view, they studied the
morphological characters of the adults and of the
development stages (eggs, larvae and nymphae) of
the five Anopheles species vector of human Plamodium they found in Algeria: Anopheles algeriensis,
A. hispaniola, A. maculipennis, A.multicolor and A.
sergenti.
They demonstrated that Anopheles was present in
all the localities where malaria occurred, confirming
in Algeria the Grassi’s aphorism: “no malaria without anophelism” (Sergent Edm. & Et., 1903b). They
described the breeding sites of these species, and
created the French term of “gîtes larvaires” for qualifying them (Sergent Edm. & Et., 1903c). They also
carried out applied investigations oriented to the
understanding of malaria epidemiology. They
defined the characteristics of the larval breeding
sites according to the species, and their seasonality.
They individualized those resulting from human
activity and developed corrective measures. They
studied the flight of the adults and their passive
transport, their feeding habits and human attraction. Lastly they studied the climatic factors related
to malaria epidemiology in Algeria.
The observations they made on the Anopheles
genus were the most complete that they developed
in medical entomology. Through long term activity
in malaria field, they acquiered a remarkable competence regarding the Anopheles species. To be
complete, I would only say that they made several
other scientific works in medical entomology, mainly in the field of transmission of pathogenic agents:
cosmopolitan relapsing fever by the human body
louse in 1908, cutaneous leishmaniasis by the phlebotomine sandfly, dromedary trypanosomiasis by
tabanids and later by stomoxes, the pigeon Haemoproteus by Lynchia maura, and lastly Theileria dispar (now T. annulata) by the tick Hyalomma mauritanicum (Dedet, 2007).
The antimalarial measures
At a time when neither insecticides nor synthetic
antimalarial drugs existed, antimalarial control measures were targeted against the human reservoir and
the mosquito vector. The global plan of antimalarial
campaign that the Sergent brothers developed
included antilarval measures, measures to avoid the
human reservoirs and individual protective measures.
The antilarval measures consisted of avoiding any
stagnant bodies of water, such as puddles, pools,
ponds, wadi, marshes. The aim was to decrease the
presence of stagnant waters, or to ensure their
drainage. The Sergent brothers summarised: “stagnant waters are unprofitable from an economic
point of view and dangerous for public health, while
domesticated running waters are useful and inocuous”. Here appears a conjunction of interest
between agricultural development and public health
improvement.
These measures generally involved large hydrolog-
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ic programmes, such as adjustment of bottom rivers,
maintenance of irrigation drains, cutting drainage
canals for drying marshes.
The alternative outflow of waters was an original
achievement of the Sergent brothers. This measure
was based on the observation that under Mediterranean climatic conditions, like those in Algeria, the
life of the Anopheline larvae lasted three weeks,
during the summer months. So, a water collection is
harmless if its duration is less than three weeks. The
idea was to change the irrigation places every week,
in order that after a week of water flow, a week of
dryness could kill the larvae. This simple measure
had a great success for land development of the
Algerian plains.
When the breeding places were not able to be suppressed, the larvae themselves were destroyed by
spreading oil or using natural predators of mosquitoes, such as the larvivore fish Gambusia holbrooki,
which the Sergent brothers have introduced into
Algeria in 1926.
The measures used to avoid the human reservoirs
were based on daily small doses of quinine, which
was administered to protect entire population (systematic quininisation). After having tested (and
discarded) the Koch method (1 g quinine every 10
days), the Sergent brothers selected the administration of a quinine daily dose of 20 to 40 centigrams in adults, as small pink pills, according to
the italian model; young children received chocolate bars containing 5 centigrams of quinine. The
distribution was made by specific persons daily,
going to private houses, the “agent quininisateur”,
who took daily note of the dose (Figure 3). The
Figure 3. Quinine systematic distribution made at private
houses by specific personnels of the Antimalaric
Department of Algeria: the “agents quininisateurs”. (Photo
from Sergent Edm. & Et., 1928).
quininisation was carried out during all the 7
months of malaria transmission. This quininisation
method had a curative effect on patients harbouring malaria parasites, and a preventive effect on
uninfected people.
The recommendation of establishing settlements
at distance of the Anopheline breeding sites and of
the reservoirs was unrealistic in a majority of the
cases, due to the impossibility of modifying the
existing situations. There was no systematic segregation programme in Algeria with separation of
the indigenous and European populations. Moreover, according to the Sergent brothers, the reservoirs included not only highly infected indigenous
population, but also the european population previously infected, even if a difference was made
between the two populations in terms of infectivity for the Anophelines, these populations being
infective respectively permanently or only during
relapses.
The antilarval measures and the systematic quininisation were completed by individual protective measures. These measures, that were the focus of the
early Algerian campaigns of the Sergent brothers,
were minimized in further campaigns. Of course
they recommended the mechanic protection of the
people by use of mosquito nets on the opening of
houses and use of bed nets. But the efficiency of
these measures was depending on the behaviour of
individuals and the care placed in nets maintenance
and use.
Some results
By the conjunction of all these control measures
regularly developed within selected anti-malarial
campaigns conducted by the Sergent brothers, a
global decrease of malarial endemicity in Algeria
was obtained. The situation varied according to the
location and the year (complete success in the
Ouled Mendil Marsh). The malaria endemicity was
reduced even if some renewed outbreaks appeared
during years with high levels of rainfall in spring,
such as the ones experienced in 1918, 1919, 1939
and 1946. The first World War was escorted by a
huge increase in malarial endemicity in all Algeria,
due to the conjunction of several factors, including
interruption of antilarval measures, dissemination
of reservoirs by troup movements, lack of quinine,
and high pluviometry in the spring of the 1917 and
1918.
The global decrease in frequency and severity of
the malarial attacks obtained in Algeria was completed by the total disappearence of blackwater
fever in the year 1928.
In light of the success of the antimalarial programme in Algeria, the Sergent brothers were asked
to develop specific antimalarial programmes in
Tunisia (1904), in the Orient Allied Armies in
Macedonia (1917), in Morocco (1919) and in Corsica (1921).
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As a conclusion
225
References
The Sergent brothers were recognised as “leaders in
malaria control”, as they were named in the British
journal Popular Science Monthly, in 1915. But they
regularly ackowledged the contribution of the Italian
Malarial School to the control of malaria. At the First
International Congress of Malaria, held in Roma in
October 1925, Edmond Sergent presented the Algerian experience through a paper on the antimalarial
methods. At the closing banquet of the same congress, he addressed the Italian organisers and
acknowledged the Italian Malarial School for its
determined contribution to malaria knowledge: “...en
ce qui concerne notre science spéciale, la Malariologie, …: si les travaux italiens n’avaient pas existé, que
serait la science du paludisme? Poser la question,
c’est la résoudre. Nous n’avons qu’à feuilleter les
publications scientifiques, nous voyons la contribution de premier ordre apportée par les travaux italiens
en étiologie, épidémiologie et surtout prophylaxie”.
The Second International Congress of Malaria was
held in Algiers in 1930, organized by the Sergent brothers. After the congress, Edmond organized field trips in
the main foci of malaria in Algeria, and all the participants attended the inauguration of the newly created village named “Laveran” as a homage to the discoverer of
the agent of malaria. Edmond was elected in 1935, in
Geneva, as chairman of the Malaria Commission of the
Hygiene Committee of the League of Nations.
Acknowledgements
The author would like to thank Yves Balard for preparation of
the iconography and Christopher Sampson for revision of the
manuscript.
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Sergent Et (1901a). Existence des Anopheles en grand nombre dans une région d’où le paludisme a disparu. Ann Inst
Pasteur 15: 811-816.
Sergent Et (1901b). Première constatation de l’existence
d’Anophèles en Afrique du Nord. Octobre 1900. C R Soc Biol
53: 857.
Theobald F (1903). A monograph of the Culicidae of the world,
III. London.
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8 DELAPORTE:Layout 1
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The discovery of the vector of Robles disease
F. Delaporte
Université de Picardie Jules Verne, Amiens, France.
Abstract. The origin and transmission of African filariasis has long remained enigmatic. Between 1915 and
1917, the pathogenic role of Onchocerca volvulus and its transmission by insects of the genus Simulium,
had been established in Guatemala by Rodolfo Robles who took opportunity of a series of discoveries to
formulate his hypothesis on the origin of Latin Americna Onchocerchiasis. The present paper gives an
historical account of the steps and the context having led to the formulation of the aetiological hypothesis and the relevant vector identification.
Key words: Robles disease, oncocerchiasis, filarial, Simulium.
In 1913, François-Marie-Frédéric Ouzilleau (18..1963), a French colonial physician working in Central Africa, wrote: “The chapter on filariasis is still
full of uncertainties. Although started well before
that of trypanosomiasis, it has not been enriched by
as much precise data as for trypanosomiasis. No
progress appears to have been made over recent
years to clarify the complexity and confusion concerning this disease; quite to the contrary” (Ouzilleau, 1913). That subject of filariasis full of uncertainties, concerned oncocerchiasis at it has been
studied in Africa. The disease was sometimes considered as a benign affection or as a disease identical
to elephantiasis. Its mode of transmission was
unknown. Three years later, a physician very rapidly
resolved all of the problems raised by onchocerciasis.
Up until then, all research on this disease had been
conducted in Africa. Strangely, the significant breakthrough was made in an unexpected place:
Guatemala. In the years 1915 to 1917, Rodolfo Robles (1878-1939)1 identified the Onchocerca volvulus
Correspondence: François Delaporte, Université de Picardie,
Faculté de Philosophie et Sciences humaines et sociales, Chemin
du Thil, 80025 Amiens Cedex 1, France, e-mail: francois
delaporte@free.fr
1 Rodolfo Robles was born in Quezaltenango, Guatemala,
in 1878. After primary school in an American college in California, he travelled to France at the age of 17 to study Medicine. He entered the “preparatory school for science university” in Rouen and got his degree in 1898. He studied medicine in Paris and graduated as a physician and surgeon of the
University of Paris. During his stay in Paris he met with E.
Brumpt in 1904. He performed several stays at the Institut
Pasteur and also travelled to the USA, Great Britain and Germany. At the end, he had got the diploma microbiology and
mycology of the Institut Pasteur. In addition to being physician and surgeon he had become specialist in hygiene and had
graduated as physician specialised in malariology of the University of Paris. He finally was member of the Geographical
Society of Washington. When back to Guatemala, he joined
the school of Medicine in San Carlos in 1905 and created a
polyclinics in Quezaltenango but failed in his attempts to create an Institut Pasteur in his town. He settled in the capital in
1910 where he became illustrious and had a number of
patients. He worked there in several hospitals, occupied several university and government positions and was elected dean
parasite and defined the clinical description of the
disease and elucidated its epidemiology.
In 1916, the Republica de Guatemala journal published the first account of the discovery of the parasite responsible for onchocerciasis in Latin America.
Robles described the sequence of events. The first
case was a young girl with chronic erysipelas of the
face associated with lymphangitis. Robles hesitated
between mechanical obstruction of lymphatic vessels
and the action of a toxin. He also formulated the
hypothesis that the filarial worm Loa loa would be
the cause of the ocular disorders, but subsequently
rejected this hypothesis in view of the negative examination. The second case was a young boy presenting
the same symptoms as the first patient. One detail
proved to be decisive in this case: excision of a
tumour on the forehead revealed a filaria similar to
O. volvulus. In the following year, Robles presented
a new version of the events in the journal La Juventud Médica (1917). When describing the first case,
he no longer mentioned the hypothesis of a parasitic
infestation related to the presence of Loa loa, but
only described the child’s periodic erysipelas accompanied by fever, warmth and pruritus of the face, due
to an unknown disease. In relation to the second
case, Robles described the ocular and cutaneous syndromes of onchocerciasis and recalled that excision
of the tumour revealed the parasite. The third version, published in the Bulletin de la Société de
Pathologie Exotique (Robles, 1919), essentially
resumed the second version: “On his forehead, the
boy had a tumor the size of a cherry which, according to his mother, had been there for several years.
When the tumor was extirpated and opened, I found
that it enclosed a fine worm, white and ball-shaped,
with the characteristics of a filaria. I understood then
that the lesions surely were due to the presence of
this parasite” (Robles, 1919a).
These accounts are historically false or, more precisely, they sin by omission, as chance and error
played a decisive role in identification of the filaria
of the Faculty of Pharmacy. Involved in public life, he has been
member of the Constituant Assembly and State Counsellor.
228
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F. Delaporte - The discovery of the vector of Robles disease
responsible for onchocerciasis. Chance, inasmuch as
this young patient spontaneously sought medical
attention and error, because Robles noted the presence of a sebaceous cyst. It was only after removal
of this cyst that he noted its fibrous consistency. The
crucial experience was therefore delayed by one
notch or one step. The surgical procedure leading to
the discovery did not correspond to removal of an
onchocerciasis nodule, but incision of a fibrous
tumour after excision of a sebaceous cyst. It is probably reasonable to say that opening of the cyst supposes its excision, but, on the other hand, it would
be untrue to say that excision of the cyst comprised
opening of the tumour and consequently discovery
of the filaria. The decisive information was published by Aguirre Velasquez, doctor and director of
the journal that published the very first report of the
discovery: “After excision of the nodule, which Dr
Robles considered to be a secondary matter, it was
the fibrous consistency of the excised tumour that
raised his curiosity. He opened the nodule with a
scalpel and was surprised to find a female filarial
worm coiled inside the nodule like a fine hair”
(Robles, 1916). That is how an apparently minor
operation can lead to a fundamental discovery.
Any persistent doubt can be eliminated by a brief
historical account. It is true that, in the history of
science, the patient’s testimony does not carry a lot
of weight, but, exceptionally, in this case, the patient
was in the best position to recount the unpredictable nature of this discovery. It is true that
demonstration of the O. volvulus filaria was purely
incidental. It is also true that chance and error are
only useful to those who know how to take advantage of them. This was the case for Robles, who, on
second thoughts and with a second cut of the
scalpel, revealed an onchocerciasis nodule when he
only expected to find a sebaceous cyst. But here is
the retrospective version of the story by the patient
himself, Ruiz Aguilar: “When the doctor performed
the first examination, he told us that he thought it
was a sebaceous growth. Several days after removal
of the lump, I returned to Robles’ clinic on Eleventh
street. To my mother’s great surprise, he told her
that my case was not as simple as it seemed. He
showed us a bottle filled with alcohol containing a
long parasite visible to the naked eye that resembled
a bit of sewing thread”2.
2
Letter to H. Figueroa Marroquin, Guatemala on 5 October 1961, pp. 66-67. A. Ruiz Aguilar is the patient in whom
Robles discovered the filaria in March 1915. The letter is
reproduced in H. Figueroa Marroquin, Enfermedad de Robles,
1963, p. 66-69. Historians often embellish the actual events;
here is an example: when Dr Robles operated on the first
oncocercoma, removing a small tumour fom the head of a
child who lived on a farm infested by the disease, he suspected the parasitic nature of this disease when he found a filamentous ball inside this tumour (Romeo de Leon J., Entomologie de la Oncocercosis, in Oncocercosis (enfermedad de
Robles), Homenaje al tercer congreso Tail-Americano de oftalmologia, Habana, enero 1948, Guatemala, 1947, p. 147).
The first difficulty encountered by Robles concerns the morphological analysis of the filaria, as the
parasite was difficult to dissect and the filaria
appeared to be sutured to the walls of the tumour.
At first sight, the large, thick cuticle and the very
obvious transverse striation suggested a filaria
belonging to the Onchocerca genus, but the absence
of a head and tail made formal identification difficult. Extraction of the whole parasite is also a delicate, long and tedious operation and must be performed by placing the filaria in water and spending
long hours carefully dissecting it. Robles invented a
new technique to overcome these difficulties, which
consisted of sacrificing dogs after making them
swallow onchocerciasis nodules. He was subsequently able to obtain whole parasites from their
stomach, which were then photographed.
Robles considered that the filaria resembled O.
volvulus previously described by Manson. The
female worm is white with a large anterior part and
an increasingly narrow posterior part. The small
mouth is followed by a clearly visible oesophagus.
The cuticle is thicker around the lips and forms a
small fold. The body presents large obvious rings
that disappear close to the posterior extremity. In
1919, Robles referred to Brumpt’s (1878-1951)
recent description of the American filaria, which
was described as being larger than O. volvulus and
differing in terms of the distribution of papillae in
the male. Actually, Robles challenged that description of the filaria found in a verminous tumor he
had given to Brumpt3. For the latter, it was a new
species, because of a differential biological feature:
the location of tumors in the scalp. A pathological
sign also differed: ocular symptoms are not
observed in African populations. “In summary, the
clear cut differences in the biological characters of
the two Oncocerques parasites of man, allows us to
differenciate these two species (…) It is beyond
doubt that the studies that will be undertaken on
the blinding Oncocerque will demonstrate the reality of that new species” (Brumpt, 1919). Robles considered that Brumpt’s description differed from his
own findings, which corresponded to the characteristics of the African filaria. The only point on which
he agreed with Brumpt was the size of the filarial
worms: the female measured about fifty centimetres,
while the male barely exceeded thirty centimetres.
It would be true to say that the most prominent
pathological signs of this local disease had been
identified for a long time, but these pathological
3
Emile Brumpt (1878-1951) has certainly been one of the
most influent French parasitologists of the first half of the
20th century. Assistant to Raphaël Blanchard, he was nominated associate professor at the Institut de médecine coloniale
de la Faculté de médecine de Paris, created in 1903 from its
beginning. He succeeded Blanchard in 1919. The Institut de
médecine coloniale was aimed at training civilian and foreign
physicians due to work in tropical areas (see Opinel A.,
2008).
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signs, as they were perceived by Guatemalan peasants clearly bear no relation to Robles disease. The
masses occurring on the head were considered to be
lumps: “The Indians do not know that the nodules
are produced by the filarial worms and, as they live
near volcanoes, they say that ‘volcano stones have
fallen on their head’ (Robles, 1919, p. 445). By calling this disease “coastal erysipelas” and “eye disease”, the Indians gave a good example of what can
be called popular knowledge, which consists of
describing the disease by its geographical distribution (the western flank of the Cordillera) and by its
most visible manifestations, such as the red colour
of the skin and eye disorders.
The pathological signs to which the Indians gave
these popular names correspond to what a
Guatemalan doctor called “myxoedema”. In 1908,
Guerrero presented the results of his survey in the
volcano region. In line with a medical tradition
solidly implanted in Latin America, he described
very distinct vast zones of goitre and myxoedema, as
the Cordillera can be divided by a horizontal line at
an altitude of about 1200 metres. Endemic goitre
and its characteristic tumours are observed above
this line, while myxoedema is observed at lower altitudes. Patients presented the typical moon face
appearance with slightly cyanotic lips, swollen eyebrows that hid the eyes, with thickened conjunctiva,
gray sclera and an opaque cornea. According to
Guerrero (1908), “the symptomatic context of
erysipelas is identical to descriptions of “pachydermic cachexia” in other countries by Raymond,
Vasquez and others: our very typical patients could
act as models for the images of myxoedema patients
published by Souques, as they present all of the
essential features”.
At the same time as Chagas described the various
forms of parasitic thyroiditis in the State of Minas
Geraes, in Brazil, Guerrero reported the existence of
thyroid dystrophy on the slopes of coastal volcanoes. Just as American trypanosomiasis must not be
confused with Chagas’ parasitic thyroiditis, it would
be wrong to consider that Robles disease corresponds to the thyroid dystrophy described by Guerrero. This is a good opportunity to raise a recurrent
question in the contemporary history of medicine:
what is the reason for this irresistible tendency to
anachronistic descriptions? This tendency simply
allows the author to show himself and his scientific
statements in a good light, illustrating that the history of science sometimes goes hand in hand with
nationalistic history.
Robles immediately described the main syndromes
of onchocerciasis. Subcutaneous nodules are typical
and are mainly situated on the head: “They are
located mainly on the head, where they tend to be
localized in the temporo-parietal region. The order
of frequency is as follows: the occipital region, then
the frontal; commonly three to four fingers distant
from the hairline; they may be present over the mastoid region and in the skin of the forehead” (Robles,
229
1919, p. 454). These nodules may be situated in the
dermis, where they are mobile when palpated with
the fingers, or in deeper regions, where they can
simulate real exostoses. By incising these deep nodules, Robles discovered these tumours that can
cause complete perforation of the skull.
The clinical features of onchocerciasis also comprise two other series of pathological manifestations: skin lesions and ocular lesions. At the acute
phase, the tense, red and swollen skin resembles
classical erysipelas of the face. These signs are
accompanied by fever. Lymphangitis results in
cracked skin giving rise to a serous exudate. The
fever resolves after several days and the patient
enters a chronic phase, characterized by indurated
oedema and eczematous, greenish pigmented, shiny
skin. The ears are swollen, deformed, and protrude
anteriorly. The eyelids are swollen and the lips are
deformed. Painful pruritus induces scratch lesions.
The eyes present particularly marked signs, usually
the pathognomonic sign of onchocerciasis, iritis, as
well as corneal ulcerations due to inflammation of
the vascular membranes of the eye, called punctate
keratitis consisting of scattered spots on an opaque
cornea. These features are associated with functional symptoms such as impaired vision and photophobia that can lead to blindness in the most severe
forms.
Robles thought that these lesions were induced by
toxins released by the subcutaneous nodules and
this hypothesis of the pathogenesis of onchocerciasis appeared to be confirmed by treatment, as excision of the nodule led to resolution of the ocular disorders. Robles reported the case of a patient in
whom excision of a solitary nodule of the hip resulted in improvement of vision and, in his young
patient, excision of the nodule was immediately followed by rapid resolution of the boy’s photophobia.
Pacheco Luna (1919) accurately summarised this
situation: “For patients with ocular disorders due to
onchocerciasis, the natural history is as follows: progressive onset of blindness in the absence of surgery
and immediate cure when the filarial tumours are
removed”.
In terms of the epidemiology of the disease, the
origin of Robles’ young patient was decisive: the boy
came from the coffee-growing region of Patulul, situated between the two volcanoes Le Fuego and Atitlan. Let’s return, for a moment, to the patient’s version of the story. At the second visit, several days
after removal of the nodule, Robles told the boy’s
mother that he would like to visit the coffee plantation: “My mother told Robles that many people on
the plantation presented the same symptoms as me
and they rubbed the stomachs of young Indians with
toads in order to cure them (although I did not
receive this treatment). During the Easter holidays,
Dr Robles, accompanied by his assistant Faustino
Gonzales Sierra, now a doctor, came to the plantation and removed a large number of nodules. They
improvised an operating table in the corridor, on the
230
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first floor for minor operations on all of the children
with nodules after shaving them (Figueroa-Marroquin, 1963)”.
In view of the large number of insects able to
transmit the disease, the first step consisted of delineating the infested zone. Robles very rapidly showed
that the zone of distribution of the parasitic infestation corresponded to a band of land between the
two volcanoes, at an altitude ranging between 600
and 1,200 metres. Two camps were situated on the
El Baul plantation: many cases were observed in the
first camp, situated at an altitude of 700 metres,
while very few cases were observed in the second
camp, situated at a slightly lower altitude, less than
600 metres. This marked disparity between two
camps situated close to each other could only be
explained by the geographical distribution of the
vector.
To verify this hypothesis, Robles started by eliminating the commonest causes of endemic diseases,
which usually correspond to two modes of contamination: contamination from person to person and
indirect contamination via drinking water. Direct
contamination had to be excluded, as the men who
picked coffee in infested regions lived in the low
altitude camp with their wives and women who had
not spent time at the high altitude camp were not
contaminated by their husbands. Men living in the
low altitude camp, although married to infected
women from the high altitude camp, were also not
contaminated. At first sight, Robles appeared to
have excluded direct contamination. However, in
reality, in the context of this parasitic infestation, he
simply showed that the vector was not present in
the low altitude camp and that it had little chance
of surviving in huts made of bamboo and palm
leaves, as they would be repelled by the smoke of
constantly lit fires.
Robles then eliminated water as a possible agent
of propagation of onchocerciasis, acting as a vector
and corresponding to the hypothesis of gastric contamination proposed for dracunculiasis and
Onchocerca gibsoni infection in cattle. As a result of
scratch lesions, intradermal worm embryos could
enter the external environment and waste water, but
embryos could also evolve like those of F. medinensis, in a crustacean that is subsequently ingested in
the drinking water. However, the river passed
through endemic zones as well as lower altitude disease-free zones. Although drinking water was taken
from various levels of the river, cases of the disease
were only observed in the high altitude camps, and
no cases were observed in the low altitude camps.
Robles presented the example of the Pentaléon and
Xata plantations situated below 500 metres, where
no cases of onchocerciasis were observed, despite
the fact that the inhabitants of these properties
drank unfiltered river water and that this river acted as a sewer for contaminated upstream properties.
Water can also be suspected to be a necessary if
not sufficient condition. Brumpt proposed the
hypothesis that the presence of water would be a
geographical determinant in the propagation of
African onchocerciasis, as the French parasitologist
thought that the disease was propagated by Tsetse
flies, especially the group composed of species with
a habitat situated close to rivers and waterholes,
such as G. palpalis and G. tachinoides. Robles
refuted this hypothesis by showing that water did
not play a role in the propagation of onchocerciasis:
“The infected farms are on the summit of a mountain or a ravine, some near a river, others very far
from water; consequently, the presence of water
seems not to have any influence, contrary to what
Brumpt says (Robles, 1919, p. 456)”.
The workers’ daily trips from one place to another suggested the possible role of black flies, as these
trips indicated the time of day of infection. Coffee
pickers worked at altitudes of about 700 or 800
metres. Infected workers living in the low altitude
camp must have contracted the disease during the
day, as they left the coffee plantations to return
home before sunset. The zone of distribution of cases of the disease in the camp situated at the same
altitude as the coffee plantations was identical to the
zone of distribution of the black fly, a small Diptera
resembling a black fly: “We think that the vectors
are two Nematocera of the Diptera family, belonging to the Simulium genus that we consider to be
Simulium samboni and the Simulium dinelli, that
currently live at an altitude of between 600 and
1,200 metres [...] They are the only known bloodsucking insects in this zone. They do not exist at
lower altitudes, although an increasing number of
blood-sucking insects are found in lower altitude
(warmer) zones”.
It is true that by exposing his experimental subjects
to black flies, Robles only added another argument
in favour of his hypothesis, without actually demonstrating this hypothesis. He did not try to prove that
the larvae of the filaria were present in the insect, or
that the insect inoculated them to man, the final
host. Robles simply established that the incriminated
species was a biting insect and that this bite was sufficiently prolonged for the black fly to absorb a drop
of blood that filled its abdomen. After biting its victim, the fly is so heavy that it can barely fly. In the
most severely affected camp, Robles exposed about
ten infested children to black fly bites at 10 o’clock
in the morning. These bare-chested children were
told not to move to avoid interfering with the flies:
“The majority of these insects rested on the ears,
cheeks, and neck; an occasional one over the forehead or thorax. Among these children there was one
with the disease in an acute stage his face was red
and swollen. For every sandfly that rested on the other chronically ill children, the child in the acute stage
was bitten by five, as if the red color attracted the
sandflies (Robles, 1919, p. 448)”.
In conclusion, Robles emphasized the fact that
women were less often affected than children and
men. His explanation for this difference was that
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contamination occurred via insect bites on the temples, neck and skull. Women were naturally better
protected from insect bites by their long hair. Robles also explained why Indians were more often
affected than Whites: not only did they pick coffee
among swarms of black flies, but they were also
more vulnerable to insect bites, as Robles had
observed that their traditional dress was a predisposing factor to contamination. Indians were the
perfect victims: they wore a shirt and cotton
trousers, leaving their neck, arms and legs exposed
and they wore a straw hat which was not sufficient
to protect the temples and the neck.
Fifteen years later, Richard Strong, professor of
Tropical medicine at Harvard University Medical
School 4, led a prestigious expedition to Guatemala.
In 1934, came out of the press his book entitled
Onchocerciasis with special reference to the central
American form of the disease (Strong, 1934).
Strong had followed Robles’ footsteps. But he did
not even mention the pioneering work that Robles
had carried out in coffee plantations.
4 Richard Pearson Strong (1872-1948). Long a member of
the professorial staff of the Harvard Medical School, he was
President of the Board for Investigation of Tropical Diseases
in the Philippines (1899-1901). He conducted researches on
many communicable diseases in many countries, including
plague in Manchuria and typhus in Serbia. He was the author
of a vast treatise on tropical medicine, conducted the course
in tropical medicine at the Army Medical School during
World War II. (Quoted from http://history.amedd.army.mil/
booksdocs/misc/evprev/ch7.htm). His name has been given to
the chair of Tropical Medicine at Harvard U. Medical School
in 1949.
Cited literature
Brumpt E (1919) Une nouvelle filaire pathogène parasite de
l’homme (Onchocerca caecutiens n sp). Bulletin de la Société de Pathologie Exotique 12: 471-472.
Figueroa Marroquin H, Enfermedad de Robles, 1963. Letter to
H Figueroa Marroquin on 5 October 1961, p 67.
Guerrero P (1908). El Bocio, el Mixedema y el Cretinismo, en
las montanas guatemaltecas. La Antigua, Tp Internacional,
231
1908, p 29. An extract of Guerrero’s text is quoted by Figueroa Marroquin, 1963, pp 76-77. The work by Robles immediately raised a controversy between supporters of the parasite theory and supporters of the myxoedema theory. For
example, see E Quintana, Un problema de semiotica nacional, La Juventud Médica, 1921, 18, pp 214-215, pp 365-366;
Reti A, Bocio, Mixedema y Filaria, idem, 1922, 19, pp 225,
471-474; Fletes SC, La Onchocerca y el Mixedema, idem,
1923, 21, pp 230-231, 551-552.
Ouzilleau F L’éléphantiasis et les filarioses dans le M’Bomou
(Haut-Oubangui). Rôle de la Filaria volvulus, Suite, Annales
d’hygiène et de médecine coloniales, 1913, 16, p 688.
Pacheco Luna R (1919). Appendice. Lésions oculaires d’après
le Dr Pancheco [sic], Bulletin de la Société de Pathologie
Exotique, 1919, 12, p 463 (461-463). See also, by the same
author, Apuntes preliminares sobre los trastornos de la vision
observados en Guatemala en los enfermos portadores de
ciertos tumores filariosos, La Juventud Medica, 1917, 17, pp
241-248; and Disturbances of vision in patients harboring
certain filarial tumours, American Journal of Ophthalmology,
Febr 1918, in Oncocercosis (enfermedad de Robles), Homenaje al tercer Congreso Pan-Americano de Oftalmologia, La
Habana, enero 1948, Guatemala, 1947, pp 41-49; and Calderon VM, Contribucion al estudio del Filarido Onchocerca
sp Dr Robles 1915 y de las enfermedades que produce,
Tesis inaugural, Guatemala, 1920.
Robles R (1916). Una enfermedad nueva en el continente ha
sido diagnosticada en Guatemala. In: La Republica de Guatemala, 29 December 1916; and in: Figueroa Marroquin H,
Enfermedad de Robles, 1963, p 60.
Robles R (1919). Onchocercose humaine au Guatemala produisant la cécité et l’érysipèle du littoral (Erisipela de la costa),
Bulletin de la Société de Pathologie Exotique, 1919, 12, p 444.
Reference of the first version: Robles R, Una enfermedad nueva en el continente ha sido diagnosticada en Guatemala, La
Republica de Guatemala, 29 December 1916; and in: Figueroa Marroquin H, Enfermedad de Robles, 1963. The journal
article, presented in the form of an interview, is reproduced on
pp 59-65. Reference of the second version: Enfermedad nueva en Guatemala, resumen de la conferencia dada por el doctor Rodolfo Robles, por Victor M Calderon, Publicado por primera vez en La Juventud Médica, Guatemala, Agosto de
1917, 17, n 8; Lecture reproduced in Oncocercosis (Enfermedad de Robles), Guatemala, 1947, pp 27-39.
Strong EP (1934). Onchocerciasis, with special reference to the
Central American form of the disease. Contribution n 6 from
the Harvard University Department of Tropical Medicine and
the Institute for Tropical Biology and Medicine, Harvard University Press.
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9 BENCHIMOL:Layout 1
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Medical and Agricultural Entomology in Brazil:
a historical approach
J.L. Benchimol
Casa de Oswaldo Cruz, Fiocruz, Rio de Janeiro, Brazil.
Abstract. Medical Entomology emerged in Brazil in the late nineteenth century, through the initiative of a
group of physicians dedicated to researching microorganisms related to diseases of public health importance, especially yellow fever and malaria. They led the institutionalization of Bacteriology and Tropical
Medicine in southeast Brazil and the sanitation of coastal cities and, subsequently, rural areas. Medical
Entomology provided the professionals who would undertake campaigns against agricultural plagues, as
well as the institutionalization of Agronomy and Veterinary Medicine. In the present article, I intend to show
how relations between the professionals who gave life to Medical Entomology in Brazil were interwoven
and to illustrate their relations with entomologists in other countries. I will also present an overview of the
research problems faced by Brazilian entomologists at the turn of the nineteenth century and early
decades of the twentieth.
Key words: Brazil, Lutz, Neiva, medical entomology, Instituto Oswaldo Cruz.
State of entomological art at the turn of the century
In March 1899, Adolpho Lutz (1855-1940), director of the Bacteriological Institute of São Paulo
(1893-1908), became an active participant in the
global survey launched by the British Natural History Museum on mosquito species that might be related to human diseases. Other Brazilian zoologists too
were engaged in this worldwide endeavour, including Carlos Moreira, of Rio de Janeiro’s National
Museum, and Emílio Goeldi, Director of Pará’s
Museum of Natural History and Ethnography1.
Having begun his studies on Culicidae in 1898,
Adolpho Lutz had material to send to the British
Museum as early as June 1899. In a letter dated
April 28, 1900, Theobald admitted that little had
been done with these mosquitoes until he had taken
the subject up two months earlier. “The whole subject is in confusion, in many cases the same insect
having been described under half a dozen different
names, simply because it came from a new locality
(…) Hence the tremendous difficulty of identification in all old descriptions – in fact, only Ficalbi’s,
Skuse’s and Arribálzaga’s are of much value”2.
Theobald had already identified two new species
of Anopheles sent by Lutz – A. albipes and A. lutzii
– and created a new genus (Aegritudines) for his river mosquitos. In 1901, Theobald published two volumes and an atlas, containing descriptions of 289
species, of which 114 were new to science. In 1903,
Correspondence: Jaime Larry Benchimol, Casa de Oswaldo
Cruz, Fiocruz, Av. Brasil 4365, Manguinhos 21045-900 Rio
de Janeiro, Brazil, e-mail: jbench@uol.com.br
1 Benchimol, 2005; Benchimol & Sá, 2007. On Goeldi, see
Sanjad (2003). Moreira, devoted mainly to agricultural entomology, still awaits a study worthy of him.
2 Rio de Janeiro National Museum Archive (henceforth
BR.MN), Adolpho Lutz Fund (FAL), folder 267. Some letters
cited in this article can be found in www.bvsalutz.coc.fiocruz.br
the first supplementary volume (III) came out, with
88 new species. Four years later (1907), the second
supplementary volume was published (vol. IV), with
a further 73 new species. Since 1903, the British
Museum had received some 12,000 specimens, of
which only about half had been examined (1907,
III-VI). The fifth supplementary volume of
Theobald’s monograph would only come out in
1910.
Theobald’s scheme for classifying different genera
was based mainly on the structure and colouring of
the scales on the insect’s head, thorax, abdomen and
wings, rather than on the size of the palpus, the feature used previously. These Diptera being ultimately
collected for their potential medical significance, it
was important to understand their life cycles and
habits, especially their proximity to human habitations and their attraction to blood.
A cursory reading of the relationship between
Lutz and Theobald might lead one to suppose it was
as one-sided as the economic ties between their
respective countries: “raw material” exported by
Lutz was converted into knowledge “manufactured”
by the British entomologist. Indeed, Theobald did
initially hold the upper hand in the relationship
because of his facilities and his access to specialized
literature and material collected from all over the
globe. The British had a broader geographical picture, which gave Theobald an advantage in comparative work. Yet Lutz had one very important advantage: he could observe the insects in their natural
habitat and handle them while still alive. He had the
chance to compare colours, movement patterns,
habitats, larvae characteristics, how they changed
on the way to adulthood and even any predatory or
peaceful co-habitation between species.
The specialist-collector relationship soon changed.
Lutz himself began to describe and classify his own
materials. He wrote a description of each specimen
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J.L. Benchimol - Medical and Agricultural Entomology in Brazil
and made observations on the habits of the imago
and larval phases of the adult specimens sent to
Theobald. Many came from larvae bred in the laboratory. Lutz always provided information on the
place where specimens were collected and whether
they were associated with the presence of humans
or not; he registered whether the mosquitoes bit
humans or other animals, whether they did this by
day, during twilight or at night, whether the bite was
painful or went unnoticed, and whether it drew
blood, emphasizing proximity or distance from
human habitations.
Lutz started to produce his own taxonomic categories with the help of what was then a scant bibliography. In his first letters, he quoted mainly Christian Rudolph Wilhelm Wiedemann (1770-1840),
George Michael James Giles (1853-1916) and Eugenio Ficalbi (1858-1922), who provided Grassi with a
systematic scheme. In their descriptions of wings,
Theobald and Lutz adopted the terminology used by
Skuse in his Monograph of the Culicidae of New
South Wales, “which is by far the simplest, and
serves for the purposes of identification perfectly”
(Theobald, 1901, VI). Another important reference
for them was Félix Lynch Arribálzaga (1854-1894).
In 1877, Arribalzaga submitted an article on mutilid
wasps – the first publication on Entomology by an
Argentine – to the Academia Argentina de Ciencias
y Letras. The following year, he published papers on
Dipterology in El naturalista argentino, the country’s
first natural science journal, established by his brother Enrique (1856-1935) and Eduardo Ladislao
Holmberg (1852-1937). In 1890, he returned to
Diptera, with two texts on Mycetophilidae. He published “Dipterologia Argentina” (Arribálzaga, 1891),
an article much used by Adolpho Lutz. Enrique’s
most important work was “Asilides argentinos”
(1879-1883) and a catalogue of the Diptera of the
Rio de la Plata river (Papavero, 1973, 335-337).
Lutz’s investigations resulted in the 1903 publication of Waldmosquito und Waldmalaria (Forest mosquitoes and forest malaria). It is therefore no surprise
that his focus was on forest “forms” collected both at
high altitudes and near rivers or the coast where
bromeliads were abundant. To undertake this programme of research, he relied on a network of collectors that included a high proportion of Swiss and
German immigrants, educated in their native countries at a time when it was commonplace to keep collections of wildlife. One of the most important species
described by Lutz is still recognized as the principal
vector of “bromeliad malaria,” which occurs in epidemic form along the coast of São Paulo state, and
endemically from São Paulo down to the state of Rio
Grande do Sul. This mosquito, which Theobald baptized Anolpheles lutzii (now A. cruzii), is also the only
known natural vector of simian malaria in the Americas (Consoli and Oliveira, 1994)3.
3
Theobald (1901, 177-178) described Anopheles lutzii. In
1905, he included it in the subgenus Kerteszia. In 1908, Dyar
Medical Entomology gains momentum
In the main, the study of disease-transmitting mosquitoes had thus far been carried out by doctors
who had acquired the skills to deal with insects on
the job, in haste, and not always most appropriately. A great deal of knowledge about culicids was
amassed but, at the same time, great confusion
arose about how to describe and name species.
Adolpho Lutz soon became the central figure for
Brazilian doctors interested in this field of research.
He supervised the first doctoral thesis on Medical
Entomology produced in Brazil. Its author, Celestino Bourroul, was born in São Paulo on 13 November 1880. In the absence of medical schools in his
state, he entered the Bahia Medical Faculty in 1899.
On the island of Itaparica, Bourroul collected mosquitoes whose habitats were bromeliad waters and
described seven species, of which one was new. A
devout catholic, he named it Megarhinus mariae” 4.
Lutz not only revised Borroul’s thesis, Mosquitos do
Brasil, published in 1904, but annexed a long paper
of his own entitled “Synopsis and systematization of
the mosquitoes of Brazil”. Lutz’s proposed grouping
of families and genera was adopted in Volume 4
(Supplement) of Theobald’s monograph – Theobald
considered it “extremely well grounded” (1907:15).
Lutz’s choice of characteristics by which to separate
the family Culicidae into two large groups was
based on whether they had a perforating or non-perforating proboscis. He further subdivided the group
with a perforating proboscis on the basis of whether
the larva had a siphon or not.
Other young Brazilian physicians were also
attracted to Medical Entomology and interacted
with Lutz. Francisco Fajardo and Oswaldo Cruz, for
example, met him during the 1893-95 cholera outbreaks in southeast Brazil. Lutz’s friendship with the
former was further enhanced by their mutual interest in malaria. Fajardo, who was also a member of
the network set up by the British Museum, earned
his doctorate at the Rio de Janeiro Medical Faculty
in 1888. He took over as assistant professor in clinical propedeutics. One of his lectures was published
as Tropical Diseases (in Port. Fajardo, 1902). In
Brazilian periodicals and in the prestigious Centralblatt für Bakteriologie, Parasitekunde und Infektionskrankheiten, he published other works that
bear witness to his interest in research on malaria,
yellow fever, cattle tick fever, among other diseases
(Fajardo, 1901). In 1893, he was elected one of the
and Knab changed its name to Anopheles cruzii, since two
species of Anopheles had already been named after Adolpho
Lutz. For more on forest malaria, see Gadelha (1994) and
Benchimol and Sá (2005, 245-457).
4
In 1913, Bourroul took over as substitute professor at the
recently opened São Paulo Medical Faculty, lecturing in
Physics and Natural History; the full professor was parasitologist Émile Brumpt. The following year, Brumpt returned to
his native France, and Bourroul took over as professor of Parasitology.
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235
youngest members of the National Academy of Medicine, with a memoir on “The malaria microbe”
(Fajardo, 1893; Benchimol, 1999). When Fajardo
visited Paris (letter dated 29 October 1900), he took
along a preparation made by Lutz from a “pernicious attack” and showed it to Laveran, who “appreciated it greatly. He authorized me to tell you that
he is willing to correspond with you and will accept
the mosquitoes that I told him you collect and
study; he is in contact with everybody” (BR, MN,
FAL, folder 180). In 1905, Fajardo gave an account
of the campaign against Stegomyia fasciata in Rio
de Janeiro at the 15th International Medical Congress in Lisbon (Fajardo, 1905). This was Fajardo’s
last mission on behalf of Microbiology and Tropical
Medicine, established under the leadership of the
group’s youngest member, Oswaldo Gonçalvez
Cruz.
In 1901, Cruz published his Contribution to the
study of culicids in Rio de Janeiro (in Port. Cruz,
1901). He had studied mosquitoes from some
malaria-infested areas near the capital. There he had
found a species of Anopheles unlike those described
by Giles: “in the absence of any pronouncement on
the subject by the elders”, stated Cruz (p. 15), “we
propose that the mosquito be given the provisional
(...) name of Anopheles lutzii, in homage to the wise
man who so skillfully runs the Bacteriological Institute of São Paulo” 5. Oswaldo Cruz graduated in
medicine in 1892 with a thesis on Water as a vehicle for microbes (in Port.). From 1896 he traveled to
France for specialized study, where he remained
until 1899. He frequented the Pasteur Institute during the boom years of the discovery of pathogenic
microorganisms, vaccines and serum therapy. He
returned to Rio de Janeiro in the year that bubonic
plague reached Santos, Brazil’s main coffee exportation port. The São Paulo and federal governments
set up laboratories to produce Yersin’s anti-plague
serum and Haffkine’s anti-plague vaccine. In late
1900, work began at the Butantã Institute, under
the directorship of Vital Brazil, Lutz’s assistant, and
at the Federal Serum Therapy Institute, created in
Rio de Janeiro. From technical director of this laboratory, also called the Manguinhos Institute, Oswaldo Cruz was appointed full Director in December
1902. Six years later, it became the Institute Oswaldo Cruz (Benchimol and Teixeira, 1993; Benchimol,
1990; Stepan, 1976).
The election of Francisco de Paula Rodrigues
Alves as president of the Brazilian Republic on 15
November 1902, brought public health activity into
the political arena. Rodrigues Alves had as his main
goal the sanitation of the federal capital, Rio de
Janiero. Oswaldo Cruz took over the General
Department for Public Health with the intention of
fighting yellow fever, smallpox, and bubonic plague
(Franco, 1969; Benchimol and Sá, 2005). The mosquito extermination squads neutralized stored water
that contained mosquito larvae. Another group used
sulphur and pyrethrum to purge houses, covering
them with huge cotton cloths to kill Stegomyia in
their winged stage. The victims of the other contagious diseases were taken with their belongings to
disinfection centres before being isolated in public
hospitals.
Meanwhile, the institute’s activities grew along
three separate lines. Biological products, research,
and teaching are still the cornerstones of the Oswaldo Cruz Foundation. Human, animal, and to a lesser extent plant diseases, defined lines of research
that put the institution in contact with different
“clients” and scientific communities. Just as the
European institutes that operated in African and
Asian colonies increasingly ventured into the field,
so Manguinhos scientists headed to Brazil’s sertão
region to study and fight diseases, especially malaria. In putting their expertise at the service of railroad companies and other private and public clients,
they faced problems different from those encountered in urban centers. They had the chance to study
little-known or unknown pathologies and to collect
biological material that greatly expanded the horizons of Tropical Medicine in Brazil (Lima, 1999;
Albuquerque et al., 1991).
5 On November 30, 1901, Lutz explained that the name A.
lutzii was already taken by Theobald, “who called one of the
two new species I sent him some time ago by that name. The
other was called A. albimanus and I fear it may be identical
to the species discovered by my colleague”. Casa de Oswaldo,
DAD, OC/COR/Ci/1901 11 19.
6 Chagas noted that after mosquitoes fed on blood, they
would get so heavy they lost their ability to fly far and stayed
inside the dwelling place while they digested the blood.
According to Chagas Filho (1993, p. 78), the importance of
this theory was only recognized at the 1925 International
Congress on Malaria held in Rome, and was only truly effec-
A new generation of versatile professionals
One of the areas that received most attention during the Manguinhos Institute’s founding years was
entomology. This enterprise was headed by Oswaldo
Cruz himself.
Carlos Chagas first contacted the institute in 1902
via Fajardo, in whose laboratory he finished his thesis, “Haematological studies in malaria” (Portuguese) (Chagas, 1903). The following year,
Oswaldo Cruz entrusted him with the task of preventing malaria in rural São Paulo, where work on
a hydroelectric dam was at a virtual standstill
because of the disease. There Chagas (1906) perfected procedures that later became commonplace
in malaria eradication campaigns. Antilarval measures were difficult in the inhospitable and uninhabited areas where hydroelectric power plants and
railroads were being constructed. The insects were
attacked mainly in their adult phase, inside homes,
where they were generally infected by patients carrying the parasite and where they themselves in turn
mostly infected healthy individuals 6.
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In 1906, Arthur Neiva joined Manguinhos. Born
in 1880 in Salvador, Bahia state 7, he graduated
from the Rio de Janeiro medical faculty in 1903. In
February 1907, he conducted a malaria campaign
in the lowlands of Rio de Janeiro state where he
proved (1910) that the recommended quinine doses were not only inadequate but also induced development of quinine-resistant strains of plasmodia. In
the following years, he and Chagas led other campaigns in inland Brazil. The development of Entomology at the Manguinhos was intimately related to
these campaigns, and the published papers of that
era focused mainly on the recognition of local
malaria transmitters. Before the creation of the
Memórias do Instituto Oswaldo Cruz in 1909, these
continued to be published in O Brazil-Medico. Neiva’s work on Myzomyia tibiamaculata (1906) was
released in this journal. “We are building a great
cage for breeding and studying the life habits of
mosquitoes, as well as transmission of malaria by
Brazilian anophelines”, commented Oswaldo Cruz
in a letter to Lutz dated 31 August 1906 (BR, MN,
FAL, folder 213).
Carlos Chagas published three papers in 1907,
which were collected in New Brazilian culicid
species (Portuguese; Chagas, 1907) 8. Lutz placed
the species described by Oswaldo Cruz in 1901
(Anopheles lutzii) in the genus Pyretophorus, created by Blanchard (1905). Theobald had then reclassified this anopheles as Myzorhynchella nigra. Chagas now asserted that “The Manguinhos wishes to
reestablish the truth of the matter, recovering the
new species (...) over which it has priority”. The
description he then provided would justify the excision of Pyretophorus lutzii and Myzorhynchella
nigra and their replacement by a “new species by
Gonçalves Cruz, Myzorhynchella lutzi” (pp. 3-4).
Chagas described two other new species very similar in appearance: Myzorhynchella parva and
Myzorhynchella nigritarsis. “The inclination of the
Manguinhos Institute”, he proudly wrote, “is to
make them varieties of the same species rather than
distinct species. However, in view of the standards
set by Prof. Theobald on this matter, we are forced
to accept the distinctive features of each anopheline
as sufficient for distinguishing the species” (p. 12).
Chagas also described Cellia braziliensis and named
a new Taeniorhynchus species juxtamansonia (Chagas, 1907). During the same period, Oswaldo Cruz
published two papers on Entomology. In the first
(Cruz 1906), he proposed a new genus in the subfamily Anofelinae – Chagasia – to include a new
tive after DDT was brought in on a large scale. In 1935 (pp.
191-231), Chagas (Chagas, 1935) wrote an exposé on malaria prevention. A vailable in Prata (1981) and at
www4.prossiga.br/Chagas/prodint/sec/pi02-318-1.html
7
For more on Neiva, see Pinto (1932), Borgmeier (1940,
pp. 1-104), Lent (1980, pp. 1581-7); Fonseca Filho (1974).
8
The book collected articles published in O Brazil-Medico
(v. 21, n. 30, pp. 291-3, Aug. 1907; v. 21, n. 31, pp. 303-5,
Aug. 1907; v. 21, n. 32, pp. 313-4, 1907).
species he named Chagasia neivae. In 1907, he proposed another genus in the same subfamily (Manguinhosia) for a species he named Manguinhosia
lutzi (Cruz, 1907). It was given the new name of
Anopheles peryassui the following year, when it was
actually found to be an anopheline. The first genus
is still valid, forming the subfamily Anophelinae,
with Anopheles and Bironella.
In a paper on malaria prevention (Chagas, 1906)
Carlos Chagas summarized the anopheles then
known in Brazil. He mentioned the new genus
recently created by Oswaldo Cruz, “with a Brazilian
species, the same that Dr. A. Lutz called Pyretophorus fajardoi. Theobald’s opinion on this matter is
expected to resolve this”. In fact, on 21 June 1906,
Cruz went into some length on the matter with
Adolpho Lutz. He had just received a specimen of
P. fajardoi from him for comparison with the Chagasia neivae. “It really is the same mosquito, but it
seems to us that we could not include it in the genus
Pyretophorus”, wrote Cruz (BR. MN. FAL, folder
213). On Lutz’s advice, he sent a specimen to
Theobald: “Mr. Lutz had described this mosquito
from a single specimen and had included it in the
genus Pyretophorus, giving it the name P. fajardoi.
But he now believes there are enough factors for a
new genus to be created”.
A picture of Chagasia fajardoi appears on the cover of The anophelines of Brazil (Portuguese), a thesis written by a student supervised by Oswaldo Cruz
and co-supervised by Adolpho Lutz. Peryassu
defended it in 1908 but did not remain at the Manguinhos Institute 9. Another recently graduated
physician who was already working there would
make important contributions to Medical Entomology and, above all, to Protozoology: In mid-1903,
Henrique de Beaurepaire Rohan Aragão went to the
Manguinhos to prepare his doctorate. He then
became responsible, among other things, for the systematic classification of ticks. This line of investigation led him to develop a vaccine for fowl spirillosis. This disease is caused by Spirillum gallinarum,
a bacterium transmitted to poultry by Argas, itself a
genus of tick from the family Ixodidae. It would
appear that Adolpho Lutz proposed the investigation because of an issue that overlapped somewhat
with malaria: Argas is the recognized vector of
Spirochaeta bacteria, but interest in it back then
was related to the discovery of Texas fever, or
bovine piroplasmosis. Today, it is known to be a
‘pathogenic complex”, which also goes under the
name cattle tick fever, affecting animals bitten by
Boophilus microplus ticks, which serve as an intermediate host for two parasites: a rickettsial of the
genus Anaplasma (Anaplasmosis) and a protozoan
of the genus Babesia (Babesiosis).
9
He took part in campaigns against yellow fever and malaria in several states. Peryassú was Director of the Pará School
of Pharmacy.
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At the beginning of the twentieth century, Texas
fever was a single entity. In 1893, Theobald Smith
and F.L. Kilborne identified its agent (Babesia
bigemina) and its transmission via a tick of the
Boophilus annulatus species. In Brazil, Fajardo
(1901) published the first paper on the disease 10.
On 16 July 1906, Adolpho Lutz told Oswaldo Cruz
that he was beginning a study of Argas and suggested that the Manguinhos take part. A week later (July
29), Cruz declared he was “entirely” at Lutz’s disposal. In September 1906, he informed the director
of the São Paulo Bacteriological Institute that Henrique Aragão would go to São Paulo to study histoplasmosis11, a disease then thought to be related to
a protozoan, and to receive further instructions on
the Argas. Aragão conducted experiments to infect
birds using both microorganisms of the genus Spirillum, and nematodes (filariae) and protozoa. In this
latter case, he followed the experimental model that
Ross used for study of the malaria plasmodium,
whose life cycle in that class of hosts had not been
entirely established.
On 15 April 1907, the Manguinhos’ director
wrote enthusiastically to Lutz: “Aragão has managed to transmit halteridium from the pigeon using
Lynchia [a genus of Diptera of the family Hippoboscidae], having verified development in the
pigeon’s lung” (BR, MN, FAL, folder 213). Aragão’s
preliminary note came out in 1907 under the title
“On the evolutionary cycle of the halteridium in
pigeons” (Portuguese; Aragão 1907). It was known
that the protozoan Haemoproteus columbae infected pigeons’ red blood cells, and its sexual reproduction had already been identified, but nothing was
known about asexual reproduction in the vertebrate
host. Aragão showed that it occurred in the pulmonary endothelium via a process called ‘schizogony’. The discovery of the exo-erythrocytic cycle of
the Haemoproteus columbae had great impact in
the centres of tropical medicine, since it helped
explain how the agents of malaria and other diseases
caused by protozoa evolved in the organisms of
their vertebrate hosts (Paraense, 1955; Fonseca Filho, 1974, pp. 42-3, 32-3).
Trypanosomes, tabanids and Englishmen
in the Amazon
In 1907, Adolpho Lutz produced a very interesting
analysis of the state-of-the-art in this field of
research. “The transmission of diseases by blood
suckers” (Portuguese) was presented at the 3rd
Latin American Medical Congress in the capital of
10 In 1888, Victor Babès had described the agent that
caused bovine hemoglobinuria (B. bovis). Anaplasmosis was
reported by Carini in 1910, in São Paulo (Carini, 1910).
11 Letter of September 27, 1906. That year, Darling had
observed the first human case of histoplasmosis in Panama
and described a protozoan. H. capsulatum was recognized to
be a fungus only in the 1930s.
237
Uruguay in 1907. In the congress proceedings,
among the presentations by Latin Americans, one
finds nothing regarding the topics dear to tropical
medicine. Lutz’s intent was clearly to instruct his
colleagues while identifying the more intriguing
unanswered questions for those who might wish to
undertake research in both human and veterinary
medicine.
In the pages related to protozoa, Lutz highlighted
the group then attracting the greatest attention in
Tropical Medicine: the trypanosomes. The first
observations on trypanosomes in Brazil had been
made by Lutz himself (1889), in rodents and batrachia. At the time of the Latin American Congress,
Brazilian investigators had their sights set on sleeping sickness, which was not found in the country. It
was transmitted by Glossina palpalis and maybe
also by G. fusca. Nagana was also transmitted to
cattle and livestock by Glossina. Around ten species
of this fly (the tsetse), which belonged to the family Muscidae, were already known. For some time,
Adolpho Lutz had been studying another family of
Diptera that seemed to be involved in the transmission of trypanosomiasis: the Tabanidae.
In mid-1907, the directors of the Institute
Soroterápico Federal and the Instituto Bacteriológico de São Paulo both set off on journeys that would
prove extremely fruitful. Oswaldo Cruz travelled to
Berlin to represent the Brazilian government at the
14th International Congress on Hygiene and
Demography; and Lutz was hired by the government of Pará to study epizootic diseases on the
island of Marajó. Lutz reached Belém in August. His
conclusions came out that same year under the title
Studies and observations about trypanosomiasis in
horses and cattle (Portuguese; Lutz, 1907; Brazil,
1907). At the end of October, in the northern Brazilian city of Manaus, he met up with a researcher
from the Liverpool School of Tropical Medicine,
Harold Howard Shearme Wolferstan Thomas, who
had just published an article on the transmission of
yellow fever to monkeys by infected Stegomyia
(1907). Two years earlier, Thomas had taken part in
the institution’s 15th overseas expedition, with a
view to setting up a laboratory in Manaus, then
under English protection. Thomas (1905a, 1905b)
had already demonstrated that atoxyl, an organic
compound containing arsenic, was efficient in treating trypanosomiasis 12. It was known that peste de
cadeira (trypanosomiasis) attacked horses in different parts of Brazil and South America, and Vital
Brazil had just published a study on the subject
(1907). The main point of reference for Lutz and
Thomas was, however, Miguel Elmassian, Director
of the Bacteriology Institute in Assuncion, who
had discovered the disease’s agent, Trypanosoma
12 For more on this, see Power (1999, 26-8, 89) and Miller
(1998, pp. 20-1). On Ehrlich’s investigations on atoxyl see
Riethmiller (1999).
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equinum. In 1903, Elmassian and Luis Enrique
Migone published an article on the subject in
Annales de l’Institut Pasteur de Paris (Elmassian
and Migone, 1903). Not only did Lutz ascertain that
the trypanosome active in Pará was the same, but he
also confirmed that capybaras were a wild reservoir
for the parasite. In fact, Lutz showed that a number
of mammals were susceptible to experimental infection (e.g., the sloth and the squirrel monkey). He
experimented with atoxyl and related substances,
but in none did he find a reliable curative agent.
Adolpho Lutz returned to São Paulo convinced that
the main transmitters of Trypanosoma equinum
were Tabanus importunus and Tabanus trilineatos,
commonly found on ranches.
Throughout his career in Entomology, most of the
new species described by Lutz were from this group
of insects. Already in 1899, he published a case of
bicheira, or myiasis of the throat, transmitted by
tabanids. When it was discovered in 1903 that
tsetse flies hosted the sleeping sickness trypanosome, his curiosity about the group was further
kindled. This led him to correspond with Etienne
Sergent following the discovery of Trypanosoma
berberum, transmitted to camels by tabanids (BR,
MN, FAL, folder 168). In 1905, Lutz published
Contributions to knowledge about Brazilian tabanids (Portuguese and German). Two years latter, Centralblatt für Bakteriologie brought out Notes on the
nomenclature and identification of Brazilian tabanids (German). The comprehensive study in which
Lutz included his observations from Pará was also
published in Germany, in 1909. That same year,
Lutz published his first two works in collaboration
with Arthur Neiva, on Tabanidae, in the inaugural
issue of Memórias do Instituto Oswaldo Cruz.(Lutz
and Neiva, 1909) 13.
The Manguinhos Institute and US entomologists
In 1908, Carlos Chagas and Belisário Pena headed
to northern Minas Gerais, where malaria was hindering construction of the new tracks of the Central
do Brasil railroad. There, Chagas’ attention was
caught by a hematophagous insect found thick on
the wattle-and-daub walls of homes: the barbeiro, or
barber bug, which has a penchant for the human
face. In March 1909, Chagas detected in the blood
of a sick child the trypanosome he had been tracking in the insect’s organism. With the aid of Manguinhos researchers, he would delve deeply into the
disease caused by Trypanosoma cruzi (Chagas Filho,
1994; Delaporte, 1999; Kropf, 2005, 2006).
Chagas’ disease consolidated Protozoology as one
of the key areas of research at the Instituto Oswaldo Cruz, while simultaneously making the Institute
an attractive place for German researchers. In July
13 All articles on Tabanidae were republished in Benchimol
and Sá (org.), 2005; all other Lutz entomological studies, in
Benchimol and Sá (org.), 2006.
1908, two professors from the Hamburg School of
Tropical Medicine came: Stanislas von Prowazek
and Gustav Giemsa. Next came Max Hartmann,
from the Institute for Infectious Diseases in Berlin.
Giemsa would return to Manguinhos in 1912, as
would Hermann Duerck, professor of Pathological
Anatomy at the University of Jena. New Brazilian
researchers joined the Instituto Oswaldo Cruz during this same period, among them Adolpho Lutz.
His relationships with universities, museums, and
research institutes certainly helped open doors for
younger colleagues who were then sent to Europe to
do specialised studies.
Arthur Neiva was the only one of these students
to go to the United States. Oswaldo Cruz made this
decision during a trip to Washington (1907-8). The
campaign against yellow fever in Rio de Janeiro had
been successful, and Cruz had just received a gold
medal in Berlin. In Washington, he met Theodore
Roosevelt and guaranteed that the US fleet could
land its crew in Rio de Janeiro without fear of yellow fever. Cruz was impressed with what he saw at
the National Museum of Natural History (Howard,
1930, p. 425). In a letter to Neiva, dated 18 July
1907, he wrote: “They’re going to topple Theobald.
They were very excited when I told them we were
completely confounded by Theobald’s orientation.
They asked me for (…) as complete a collection as
possible of our mosquitoes, most of which they do
not know” (Fundação Getúlio Vargas, Arquivo
Arthur Neiva, ANc May 3, 1925).
At a crucial historical moment, the Manguinhos
was thus strengthening its ties with another community in entomological research, and one that was
about to cause a major upset among those who had
until then been deemed the undisputed authorities
in this field. This choice was also influenced by a
factor of ecological importance: South and North
Americans alike needed to investigate neo-tropical
fauna since the British had only indirect access to
these species. Working in loco, the Americans were
able to cooperate (or compete) in their efforts to
compile a more extensive inventory and better to
observe interrelations between groups and their
environments. Much as in Europe, agricultural pests
stimulated entomological studies in the US
(Howard, 1930; Mallis, 1971). In 1881, the US
Department of Agriculture established an Entomology Division. Its recruitment included Daniel
William Coquillett, who became honorary curator of
the Diptera section of the NMNH in 1896. His book
on The type Species of North American Genera of
Diptera (Coquillett, 1910) was received with great
acclaim by the entomological community. In all,
Coquillett described some one thousand species of
the group. He maintained a steady exchange of ideas
with Leland O. Howard (1857-1950) during the latter’s time as head of the Entomology Division.
Interest in Diptera rose sharply after Ross and the
Italians discovered their role in malaria and the North
Americans in Cuba had proved that the insects
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spread yellow fever. Howard, who had already studied the biology of Culex quinquefasciatus, published
a paper on Anopheles quadrimaculatus, the country’s
main malaria vector. In 1901, he released Mosquitoes: How they live; How they Carry Disease; How
they are Classified; How they may be Destroyed
(Howard, 1901). In 1902, Howard requested funding
from the Carnegie Institution to study American
Diptera. He contended that the work of Theobald
and Giles did not contain material representative of
North and Central America and the Caribbean. When
funding was made available in 1903 (Howard, Dyar
and Knab, 1912), Howard invited Frederick Knab
and Harrison Gray Dyar to collaborate in this ambitious venture (Dyar, 1905, 1906; Dyar and Knab,
1906). Knab had had the opportunity to develop his
entomological background while on an expedition
down the Amazon River in 1885-6. He joined the
Department of Agriculture’s Entomology Division in
1906. Following Coquillett’s death (7 July 1911), he
became curator of the Diptera collection at the
NMNH 14. Harrison Gray Dyar had been working
there since 1897, as head of the Lepidoptera section.
His research on larvae produced Dyar’s rule, which
established the insect’s developmental stage by measuring the size of its head. Dyar and Knab were
responsible for the taxonomic part of the work organized by Howard. In addition to writing many articles
about North American Lepidoptera, Dyar investigated mosquitoes, especially in their larval stage. His
research on the male genitalia was vital to classification of the group. The Mosquitoes of North and Central America and the West Indies (Howard, Dyar and
Knad, 1912; 4 volumes published between 1912 and
1917) was a landmark in the taxonomy of Diptera
and sealed a long-running dispute over taxonomic
norms.
Just as Theobald’s monograph played a role in the
construction of the British Empire, the undertaking
by Howard and his collaborators was instrumental
to the expansion of US imperialism. In 1901, President Theodore Roosevelt announced his Big Stick
doctrine, corollary of the Monroe Doctrine (1823),
whose slogan had been “America for the Americans”. Its first fruit was the separatist movement in
northern Colombia, fomented by the US, which ultimately resulted in the 1903 creation of an independent state on the Isthmus of Panama. Construction
of a canal to link the two oceans began the following year. World War I ultimately cleared the way for
the US to take over markets and territories controlled by Britain and to expand its influence beyond
the Caribbean and Central America.
Adolpho Lutz, also an authority to reckon with
As noted above, Lutz published a new taxonomic
scheme of Culicidae as part of Celestino Bourroul’s
14
He passed away in Washington, on November 2, 1918,
victim of an undiagnosed illness he contracted in Brazil.
239
thesis in 1904; he grouped genera into different subfamilies, making this suprageneric division using larval characters for the first time. Lutz’s scheme
gained international attention thanks to Raphael
Blanchard, who dominated the network of zoologists and parasitologists around the world. In 1905,
in Les moustiques: histoire naturelle et médicale, he
reproduced the classification proposed by Lutz (pp.
619-20)15. It earned praise in the US, especially
from Dyar, who was working with larvae and their
genitalia. In 1906 (p. 188), he remarked:
“A classification proposed by Dr. Lutz and cited in R.
Blanchard’s work (1905) corresponds precisely to larval characters, and this is obviously the best and most
natural classification proposed to date. Dr. Lutz
achieved this felicitous result not by using any new
character but by altering the order of importance of
the old ones: the relative length of the palpi in the
male or female. Up until now viewed as the prime
character, it has been relegated to a secondary plane
(…) The useless character of scales, used by
Theobald, has been discarded, and rightfully so. I am
referring to primary divisions, or subfamilies, without
entering into the merit of the classification of genera”.
The critique of the characters used by Theobald and
other researchers, including Blanchard, for the separation of subfamilies and genera would be further
developed in an article published by Dyar in collaboration with Frederick Knab (1906, pp. 169-230).
“That larval characters are of great value and interest there is no doubt” – wrote Theobald (1907, pp.
9 and 13) – “but to form genera and species on larvae is surely unusual”. Whoever examined a broad
series of any larvae, he argued, would note much
variation in their characters, not only in different
stages of the same species but also in the same stage
across different specimens of the same species.
Theobald made reference to another classification
method proposed by E.P. Felt (Felt, 1904), which
took into account not only larval characters but also
the male genitalia and nervure of the wings. Dyar
(1905, pp. 42-9) agreed with Felt and concluded
that genital divisions were corroborated by larvae,
thereby constituting a more natural division than
that based on scales and palpi. In the 1907 volume,
Theobald reiterated the value of the structure of
scales for diagnosing specimens, but he adopted
Lutz’s taxonomic arrangement with minor changes.
Thereafter their correspondence dwindled. Theobald
concentrated more and more on his first area of
interest – agricultural pests – while Lutz began his
new life at the Manguinhos. Just as did the British
entomologist, Lutz now relegated mosquitoes to a
secondary plane among the entomological groups to
which he devoted his time. He turned to the Simuliidae, first and foremost, followed by Phlebotomus
15 The two began corresponding in 1901 or perhaps even
earlier. In June 1905, Blanchard requested specimens of mosquitoes for his entomological collection, “qui ne comptait
pour ainsi dire aucun type sud-américain” (BR, MN, FAL,
folder 255).
240
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flies, Ceratopogoninae, Megarhininae (on the eve of
World War I), Hippoboscidae, Oestridae, Trypaneidae and, lastly, Blephariceridae. At that time, these
groups had not yet been associated with any notable
medical or sanitary issue, with the exceptions of the
Tabanidae, which Lutz studied until his death, and
Phlebotomus flies, associated with leishmaniosis.
Arthur Neiva in the United States
In April 1910, Arthur Neiva arrived in Washington.
Howard (1930, p. 425) would later describe him as
“primarily perhaps a bacteriologist, but tremendously interested in Medical Entomology”. On 11 July
1910, Neiva wrote Lutz: “I’ve read almost all of
Dyar and Knab’s [book], and I am certain that
although quite revolutionary, it will eventually prevail (…) This is proving to be Theobald’s ‘Way of
the Cross’: he’s being hit left and right; it makes you
feel sorry for him”.
In this letter, Neiva analyzed the state of the art in
relation to different groups of Diptera: “excepting
Culicidae, it seems everything is yet to be done. “His
main reference was Samuel Wendell Williston’s
Manual of North American Diptera (1908). There
he had found a didactic presentation of Dyar and
Knab’s old classification of mosquitoes, along with
those proposed by Schiner, Coquillett and other
entomologists. Neiva noted their shortcomings:
Williston’s account of the simuliids was rather weak;
and Hine had studied tabanids, but “it is nothing
remarkable; he adopts only one family: Tabanidae”.
While in Washington, Neiva’s attention was captured by Megarhinus16. He realized that the Brazilians had a much broader understanding of this
group of mosquitoes thanks to their observation of
larvae and egg-laying.
We are exceptionally well positioned right now (…)
They do not concern themselves with this mosquito in
their monograph (...) As you know, they have separated out some species of it, and I believe they have
made a muddle of things. (…) Most Brazilian
Megarhinus are not known here, meaning that since
we have the two largest known collections available,
we can compile a decisive study. They are also convinced here that Meg do not feed on blood (…). Don’t
you think this is a wonderful opportunity, with the
possibility of revising Megarhinus from around the
world? (BR, MN, FAL, ibidem).
In 1913, Lutz and Neiva published “Contributions
to the biology of Megarhininae with a description of
two new species” (in Portuguese and German). They
studied the habits and habitat of these species, from
egg-laying to larva to adulthood. The following year
saw publication of the paper on the species
Megarhinus haemorrhoidalis. The authors presented an extensive synonymy, based on material examined by Neiva in the US capital and then in Europe.
Neiva made a sufficiently good an impression on
16 In 1908, Neiva described the species Megarhinus fluminensis in Peryassú’s dissertation (Theobald, 1910, p. 90).
Howard that he was invited to write for The mosquitoes of North and Central America and the West
Indies. Neiva’s text was published in volume 1 (pp.
188-94) under the title “The Malarial Organisms”.
Here he analyzed existing types of plasmodia and
their life cycles. At the suprageneric level, he adopted the new order Binucleata, created by Max Hartmann and Victor Jollos. As mentioned earlier, Hartmann had been at the Manguinhos in 1909, at the
climax of Carlos Chagas’ discovery, and he worked
with him on a number of questions in Protozoology
(Sá, 2005). Regarding classification at the specific
level (genera and species), Neiva followed Blanchard.
Neiva’s studies in the US encompassed other
groups of insects, especially Hemiptera of the genus
Triatoma, whose medical importance had just been
revealed by Chagas. Neiva had analyzed the biology
of Conorhinus megistus Burm in a paper published
in 1910. Writing to Lutz on 26 October, he commented: “If Dr. Oswaldo wishes, Manguinhos could
take the lead in these studies, for they are neglected
everywhere”. At the end of 1910, he visited museums of natural history in Paris, London, Vienna,
Berlin, and Copenhagen. Between 1911 and 1913,
he was to publish descriptions of new species from
the African continent, South America and the US
and, lastly, in 1914, he would complete Revision of
the genus Triatoma (in Portuguese), which received
an honorable mention from Rio’s Medical Faculty,
where Neiva was appointed professor in Medical
Natural History and Parasitology.
In his last letter from the US, dated 26 October
1910, Neiva mentioned Lutz’s second work on
simuliid (blackflies), which would be published later that same year. In his first communication on the
topic (1909), Lutz had described eleven species
found in Brazil, of which five were new and one a
new variety unique to the region. In addition to
proposing a taxonomic key for these species, he analyzed their biology, ecology, and physiology and provided a detailed explanation on how to raise their
larvae and pupae in the laboratory. In his second
communication, the number of known species rose
significantly. In light of the “new orientation” (1909,
p. 214), Lutz created a taxonomic key based on the
pupa stage to determine the species of Simulium.
Simuliidae would be the object of one more article,
in 1917. In Knab’s opinion (1911, pp. 172-9), these
papers constituted the most complete study ever
produced on this group of Diptera: “Dr. Lutz is not
an old-school systematizer, for he addresses his topic from all angles. He places full value on data
obtained from the initial stages and through biology,
relates this to the imago characters, and, at the same
time, carefully takes into account possible sources of
error”.
Lutz had already called attention to the morphological changes occurring in the designs on the
shields of simuliids, depending on the impact of
light and the conservational state of the specimens
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– an observation important in identifying specimens
stored in scientific collections. Furthermore, he
recorded changes in the body colourings of these
Diptera caused by pigments deposited in their tissues following hemolysis of the hemoglobin in the
blood ingested by females (Amaral-Calvão and
Maia-Herzog, 2003, p. 263).
Other components of the entomologist’s network
One of Lutz’s closest collaborators up to 1915, Neiva developed the discipline in new directions as he
founded and directed scientific and public health
institutions17. Both his experience and accumulated
bibliography were fundamental to studies on
Diptera at the Manguinhos. Just as important a role
was played by the increased exchange with North
American and European institutions. The Italian
Mario Bezzi (1868-1927) and the German Paul
Speiser were among the entomologists who traded
specimens and knowledge with Lutz and Neiva.
In that year the long-running correspondence with
Joseph Francisco Zikán began. Born on 1 March
1881, in Bohemia (then part of the Austro-Hungarian Empire), Zikán emigrated to Brazil in 1902. He
worked first at a foundry in São Paulo, and then as
an elementary school teacher in Minas Gerais. Interested in collecting butterflies from childhood, he
managed to reconcile his butterfly-catching activities
with those of an administrator of rural properties.
At Brazil’s Centennial Exhibit in 1922, his insect
collection received a prize (Nomura, 1997, pp. 812). Hired as a technical assistant at the Biological
Station (now the Itatiaia National Park), Zikán
made a decisive contribution to the cataloguing of
the numerous species found in the region18.
In 1909, Lutz began his correspondence with
Charles Townsend, a member of the Office of Entomology of the US Department of Agriculture.
Townsend had been hired, in 1910, by the Ministério de Fomento in Peru to create its entomological service. On 25 January 1913, he informed Lutz
that he had begun investigating the transmission of
Oroya fever and asked for works on South American hematophagous Diptera. He believed this disease to be transmitted by ticks and so he was par-
17 That same year, he was hired to create a division devoted
to medical zoology and parasitology within the Buenos Aires
Bacteriological Institute. In December 1916 the São Paulo state government invited him to head its Serviço Sanitário
(1917-8). In January 1923, he became head of the Museu
Nacional do Rio de Janeiro and the following year led the
campaign against the coffee borer in São Paulo, which in
1927 resulted in the creation of the Instituto Biológico de
Defesa Agrícola e Animal. On this topie, see Silva (2006).
18
In 1952, Carlos Alberto Seabra, a rich amateur entomologist, bought Zikán’s entomological collection for Manguinhos Institute. Zikán published some sixty papers on several
groups. In 1940, together with his son Walter, he began a catalogue of insects from the Mantiqueira Highlands. He passed
away in São Paulo city on May 23, 1949.
241
ticularly interested in Aragão’s article on the Ixodidae of Brazil (Aragão, 1911). On 27 May, Lutz suggested to Townsend that phlebotomines and ceratopogonids were the more probable transmitters of
the disease. The papers on phlebotomines received
from Marett and Newstead, and further research,
led the US entomologist to confirm this hypothesis
and to ascertain that native dogs were susceptible to
the natural infection19.
Another of Lutz’s interlocutors who deserves mention is Hermann Friedrich Albrecht von Ihering,
head of the Museu Paulista since 1894, and founder
of Revista do Museu Paulista, to which he was one
of the main contributors. On September 13, 1909
von Ihering informed Lutz that he had been charged
with organizing the committee to represent Brazil at
the First International Congress of Entomology.
Ihering asked Lutz to provide him with a “list of the
names of worthy entomologists” (BR, MN, FAL,
folder 157). The Congress took place in Brussels in
1910, and was chaired by Belgium scientist Auguste
Lameere (1864-1942). Most participants were delegates from Europe. Of the entomologists who supported the idea in other continents, few attended.
The only paper from Brazil was presented by
Walther Horn, from Berlin, and had been written by
Zikán; it concerned larvae of the family Cicindelidae
(1er Congrès international, vol. I, pp. 69-84).
Medical Entomology at Instituto Oswaldo Cruz
in the 1910s and 1920s
With a growing number of groups represented in its
entomological collection, the Institute Oswaldo
Cruz began to play a role similar to that of a national museum of natural history (see Sá, 2008). The
collections grew markedly during the 1910s, thanks
in part to medical-sanitary expeditions to the interior of Brazil at the behest of private companies and
federal government agencies (Albuquerque et al.,
1991; Lima, 1999).
In 1910, the Madeira-Mamoré Railway Company
hired Oswaldo Cruz himself. Initials work had been
interrupted but was recommenced in 1907 by a
company put together by US entrepreneur Percival
Farquhar. Before his departure in a debilitated state
of health, Dr. Belt, head of company’s medical team,
warned: “The region to be crossed [...] is the most
disease-ridden in the world” (Ferreira, n.d.). On 16
July, Oswaldo Cruz travelled to Porto Velho (Roráima). In his report to the company, Cruz (1910,
1913) described the nosological situation in this
region, where beriberi and pneumonia manifested
themselves in extremely severe forms; and malaria
attacked 80% to 90% of the people. In 1912-1913,
a team led by Carlos Chagas traversed a large part
of the Amazon River basin (Cruz, 1913), Arthur
19 BR, MN, FAL, folder 157. In 1919, he was hired by the
São Paulo government to work with insects injurious to agriculture. On Oroya fever, see Cueto (1992).
242
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Neiva and Belisário Penna covered 7,000 kilometres
in the states of Bahia, Pernambuco, Piauí, and Goiás
(Penna and Neiva, 1916), and Adolpho Lutz sailed
the São Francisco River and some of its branches
(Lutz and Machado, 1915).
In 1911, Lutz and Neiva described two new
species of Culicidae (Culex scutipunctatus and
Anopheles matogrossensis), one found in northeastern São Paulo and the other in Mato Grosso. In
1912, they published a most interesting study on a
parasitic fly that feeds on birds, Mydaea pici
[Philornis]. Their paper on Phlebotomus also came
out in 1912. Although the role of Phlebotomus as a
disease transmitter was still unknown, the authors
stressed the voracity with which the females
attacked humans and fed repeatedly on their blood;
indeed Lutz and Neiva speculated that their role as
transmitters “of certain illnesses seems at times certain, at times very likely” (Lutz and Neiva, 1912, p.
84).
There had in fact been speculations about the relation between phlebotomines and leishmaniasis since
1909 (Dedet, 2005). Adolpho Lutz also advanced
this hypothesis in the midst of an interesting controversy with Frederick Knab (Benchimol and Sá,
2005, pp. 148-9). It began with an article in which
Knab (1912, pp. 196-200) analyzed the transmission of diseases by bloodsucking insects. Only
insects closely associated with man, he wrote, and
which regularly sucked human blood, could host
and transmit a parasite found there. The argument
contradicted what Lutz had posited in the article on
forest malaria (1903). For Knab, the Anopheles
incriminated by Lutz probably had nothing to do
with the outbreak of malaria among the workers
camped in the Santos highlands. When they arrived
there, Knab believed, these men brought latent
malaria with them, and work-related problems
caused the disease to manifest itself.
Adolpho Lutz argued that two transmitters of
malaria in Brazil – Cellia albimana and, mainly, Cellia argyrotarsis – were often found in uninhabited
places. Men who ventured into areas where large
animals were rarely found of course attracted mosquitoes, and if they stayed there long enough: “the
epidemic [would] accompany the growth of the
infection among the mosquitoes, and they themselves [would] grow in numbers thanks to the easy
feeding. It is a well-established fact that a species
can become an excellent intermediate or definitive
host of a parasite new to a region because the host
for the following stage was only recently introduced
(1913, 108-9)”.
Lutz was already, it seems, envisaging the possibility that humans could be involved in existing or
emerging forest cycles, and not just in the case of
malaria. In a later communication (Lutz, 1913b), he
stated:
Misters Dyar and Knab think mosquitoes that have
never been in contact with men cannot transmit disease (…) but it so happens that in Brazil roads and
railways have been built under such conditions, and
nearly always there have been malaria epidemics. I
know also of epidemics of Leishmania sores, with
good reasons attributed to transmission by Phlebotomus, observed in absolutely deserted zones. I have
also seen a small yellow-fever epidemic amongst people living in a place where wood mosquitoes could be
expected (…) All that is missing is that the transmitter, whatever its past may be, belong to a category in
which the parasite can thrive; then, it must have
repeated access to human beings, some of them
infected and some lacking immunity. As the process of
development takes time, its life must not be too short.
For that reason, oviparity is a favorable condition.
A few months later, on 14 July 1914, Cruz reported
to Neiva a communication by Sergent (probably Etienne) at the Société de Pathologie Exotique, in Paris.
He wrote: “He said he found that in areas with leishmaniasis there are also large quantities of Phlebotomus (...) He raises the hypothesis that the Phlebotomus is the transmitter (here you are well ahead) and
the gekko, the depositary of the virus. We are in
favourable conditions to verify the fact (...) You could
very well take care of this and prevent us from a new
defeat like the one in Bauru” (FGV/CPDOC, Archive
Arthur Neiva, ANc 03.05.25).
Cruz referred to the Bauru ulcer, which spread
through São Paulo state during construction of the
Nordeste Railroad. The cutaneous and nasopharyngeal lesions observed in individuals working in the
forest hinterlands were confirmed, in 1909, as
deriving from leishmaniasis. This finding was made
almost simultaneously by Adolfo Lindenberg, at the
Bacteriological Institute, and by Carini and Paranhos, at the Institute Pasteur, both in São Paulo.
In 1911, at Instituto Oswaldo Cruz, Gaspar de
Oliveira Viana classified the Bauru ulcer and the
Amazonian ulcera brava as leishmaniases, and
described a new species, Leishmania brasiliensis, as
the agent of the leishmaniasis observed in several
regions of Brazil and Latin America. Lindenberg
had named the disease “ulcerous leishmaniasis.”
(Brumpt and Pedroso, 1913) and Pedroso, in 1913,
proposed the term “American leishmaniasis of the
forests,” emphasizing the supposed ecology of the
disease. The dermatologist Eduardo Rabello (1925)
preferred “tegumentar leishmaniasis” to distinguish
it from visceral leishmaniasis, considered non-existent in Brazil. In Argentina, Neiva (1917) found
Leishmania brasiliensis and showed that it dated to
pre-Colombian times – its lesions were depicted in
Inca ceramics. Neiva also argued that the Phlebotomus were transmitters of tegumentar leishmaniasis.
In 1921, the Sergent brothers and their co-authors
L. Parrot, A. Donatien and M. Béguet (1921) confirmed the role of Phlebotomus in the transmission
of cutaneous leishmaniasis (also called Oriental sore
or Biskra button). A year later, Henrique Aragão
(1922) reported that one of the species described by
Neiva and Lutz, Phlebotomus intermedius (now
Lutzomyia [Nyssomyia] intermedia) was the vector
of Leishmania (Viannia) braziliensis. Aragão relat-
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ed the density of these Phlebotomus to the incidence
of tegumentar leishmaniasis in Laranjeiras Valley,
Rio de Janeiro city. In his experiments with dogs,
ulcers containing amastigote formats of the protozoan were produced (Rangel and Lainson, 2003).
Another species described by Lutz and Neiva,
Phlebotomus longipalpis (now Lutzomyia longipalpis), was associated to the American visceral
leishmaniasis (Calazar) by Evandro Chagas (19051940), son of Carlos Chagas, in 1936. This finding
was due to observations made by Henrique Penna
two years earlier, when working for the Rockefeller
Foundation’s Yellow Fever Services. The analysis of
liver fragments removed in viscerotomy posts in
Northeast Brazil showed that 41 deaths were due to
visceral leishmaniasis (Brazil-Medico, n. 48, 1934,
pp. 949-50). A team led by Evandro Chagas studied
the disease in Northern Brazil and in Argentina, and
published a report the following year (Chagas, Cunha, Oliveira Castro, Castro Ferreira, and Romaña,
1937, pp. 321-90).
Lutz had other collaborators besides Neiva at the
Institute Oswaldo Cruz: Gustavo Mendes de
Oliveira Castro, with whom Lutz studied tabanids,
and Ângelo Moreira da Costa Lima, who would later (1939-1962) devote himself to the monumental,
12-volume Insects of Brazil (in Portuguese). These
physicians belonged to a generation that reached its
professional maturity at a moment of greater opportunities in the training and practice of Entomology
as a specialty, both in Medical Zoology (rural sanitation programs) and in Agricultural and Veterinary
Medicine.
In the 1920s, a plague threatened coffee, Brazil’s
main agricultural export crop. Neiva and Costa Lima
were able to identify Stephanoderes hampei, an
agent that had caused great devastation in Java and
Sumatra, compelling many regions to substitute rubber plantations for coffee. A commission led by Neiva started the campaign against the plague, combining research, inspection, and educational work. At
the end of 1927, the commission was replaced by a
permanent organization, the Biological Institute for
Agricultural and Animal Defense, still under Neiva’s
leadership. Based on models derived from Public
Health, the campaign was a landmark in the institutionalization of agricultural research in Brazil and in
the pioneering use of biological control against plant
plagues (Silva, 2006).
In 1928, yellow fever struck Rio de Janeiro again.
The infection of rhesus monkeys in French West
Africa that same year undermined existing animal
models and etiological theories. The theory of
“exclusive” transmission by Aedes aegypti, as sustained by Oswaldo Cruz and the idealizors of the
campaign launched by the Rockefeller Foundation
immediately after WWI, unravelled at Canãa Valley
in rural Espírito Santo in 1932. A study by Soper
and collaborators (1933) proved that yellow fever
was spreading endemically in the interior of Brazil
and that it had an unknown number of hosts and
243
vectors. A new cycle of zoological research was initiated, with wide collaboration between investigators in the Americas and Africa, prompting interactions between entomologists, sanitarians, and
researchers in the emerging field of Virology (Benchimol, 2001; Lowy, 2001).
Conclusion
The global effort regarding what are now known as
emerging and re-emerging diseases would have
failed had it not been for the painstaking work of
Adolpho Lutz and other pioneers of medical entomology. The course of their careers was determined
by the great public health challenges before them
and by clashes between nations vying for world
domination, in which, as we have seen, there was a
direct correlation between the agendas of the entomologists and those of the established and emerging
empires. Great Britain and the US appear to have
dominated entomological research. In both countries, it was undertaken largely by professionals and
institutions that had amassed considerable experience in what was called “Economic” Entomology –
the branch of the discipline that dealt with agricultural pests.
Although some work had been done on this subject in Brazil, mainly at the Museu Nacional do Rio
de Janeiro, entomology relted to the investigation of
human and animal pathologies only really developed
once doctors with expertise in the study of bacteria,
protozoa and other pathogenic parasites became
involved. The Bacteriological Institute of São Paulo
and the Manguinhos Institute stand out as the pioneering institutions of Brazilian Medical Entomology. The discipline gained new momentum in the
1930s, when professionals graduating from Medical
Faculties found sufficient employment in teaching,
fieldwork and research exclusively within the specialty. And it was doctors skilled in the study of disease-transmitting insects – Arthur Neiva and Costa
Lima, to name but two – who were instrumental in
strengthening Agricultural Entomology and in rescuing and renewing research traditions in this area
from stagnation among the museums of natural history.
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Insects variation and
adaptation to environment
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Culicoides and the Tartar Steppe: Il Deserto dei Tartari
Culicoides and the spread of blue tongue virus
R. Houin
Professeur Ecole Pasteur - CNAM de Santé publique, Paris, France.
Abstract. Culicoides were described for the first time in England in 1713, but named by Latreille in 1809
only. Even so, they were better known as Ceratopogon until Kieffer reintroduced the name Culicoides. The
family name became Ceratopogonidae, the description by Meigen (1803) being better adapted to that
systematic level. Culicoides were considered simply as biting insects until it was found that they can carry filaria and viruses. In 1944, du Toit in Transvaal described their role in the transmission of blue-tongue
virus. Blue-tongue disease has since extended progressively northward from South Africa, disseminated
by Culicoides imicola. At the end of the 20th century, it reached the southern shores of the Mediterranean
sea, and has since threatened the southern Europe. Surveillance and prevention procedures were put in
place, but fortress Europe was taken breached when a different strain of the virus entered through Belgium in 2006. Transmitted by local Culicoides species that were aggressive and abundant, the disease
spread quickly, in a disastrous epizootic southward through more than half of France. Westward, infected
insects have been carried by wind over the Channel, introducing the disease to England.
Key words: Culicoides, blue tongue fever, midges, Ceratopogonidae.
Culicoides are tiny insects, usually known as biting
midges. They would have remained unstudied for
decades (even centuries!) among the world of gnats,
had they not administered painful bites to animals
and man. This unpleasant characteristic would not
have been sufficient to bring them to prominence
had they did not suddenly revealed themselves,
three centuries after their first description, as the
agents of an animal plague: blue-tongue disease,
which threatens sheep (but also cattle) farming
throughout Europe. This epizootic outbreak was
preceded by a slow northward progression of the
disease from its historic South African heartland.
The contrast between these two patterns of spread,
and the consequences of the recent drastic change
of habit, are certainly worth highlighting. The
spread of blue-tongue virus is taken here as an
example of the consequences that variations in a
virus on the one hand, and associated changes in the
insect vector on the other, can have on the timing
and course of an epizootic.
The evolution of Culicoides taxonomy
In the 18th century, the painful bites of Culicoides
led early to their precise description. In 1713, the
Reverend Derham, in England, published a work
entitled Physico theology: or a demonstration of the
being and Attributes of God from his works of creation in which he described a small midge which he
called Culex minimus nigricans, maculatus, sanguisuga. He also reported that they were called nidiol in the country of Essex. He linked the adult insect
Correspondence: René Houin, 6 rue Coëtlogon, 75006 Paris,
France, Tel 33 1 45483015, e-mail: houin.rene@neuf.fr
with larvae found in swamps, and specified that, in
some occasions, the adults could fly in their thousands near streams of running water. He insisted on
the harmfulness of these creatures to both man and
animals, quoting the example of horses spotted all
over with blood following attack by a cloud of these
creatures. Some years later, in 1758, Linnaeus
described Culex pulicaris in his seminal work, Systema naturae. Even if Derham’s description was not
in accordance with the criteria of systematic typing,
it is generally recognised that the description given
by Linnaeus deals with the same insect.
Some fifty years later, in Paris in 1809, Latreille
esteablished the genus Culicoides and defined it in
his book, Genera crustaceorum et insectorum.
Although the description was excellent, most 19thcentury entomologists preferred the name Ceratopogon created by Meigen in 1803. For this reason,
several species of Culicoides were described initially as Ceratopogon. It was only at the begining of the
20th century that it was recognised that the
Meigen’s description included more than one genus,
and was revised as that of the Ceratopogonidae family. Ed. Sergent, in Les Insectes piqueurs et suceurs,
published in 1909, did not mention any genus name,
but only the family. Long after Derham, another
clergyman, J.J. Kieffer played a major role in the definition of midges. In 1911-1912, he reintroduced
the genus created by Latreille and included it in the
Ceratopogonidae family. At that time, these insects
were considered no more than a nuisance, which
probably explains why Raphael Blanchard did not
include them in his Traité de zoologie médicale
(1890). Veterinarians were more interested, as indicated by L. Gedoelst in his Synopsis de Parasitologie de l’homme et des animaux domestiques, published in Brussels in 1891. This contained a draw-
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R. Houin - Culicoides and the Tartar Steppe
ing, together with a description and a list of 7
species, 4 of them of African origin. In 1917, J.R.
Malloch, describing some species from North America, brought the family close to the Chironomidae,
as it is today, within the superfamily of the Chironomoidea. Some further genera were added,
including Leptoconops, another fierce biter. Later
still, new species of Culicoides were described from
all around the world. There were quickly so many
(now about 1,400) that it became necessary to collect the species within the genus into morphologically defined clusters, a task which was achieved by
J.W.S. Macfie in 1940. Of special importance to
what follows, are the works of B. De Meillon,
O.G.H. Fiedler and J. Clastrier in describing the
most important African species, among them Culicoides imicola.
Culicoides as vectors of filariosis
The ability of certain species of Culicoides to transmit infectious diseases has only recently been
proven. The first parasitic organism discovered to
be transmitted by culicoides, reported by N.D.A.
Sharp in 1928, was a filaria, Dipetalonema perstans, common in some African countries, such as
Cameroon, where 92% of the human population
carried the larval form of the parasite in their blood.
Yet Sharp found only 7% of the midges to be infected. He established that a week was enough to
ensure the development cycle of the worm in the
insect, which becomes infectious upon biting. However, this filaria was then considered harmless. J.J.C.
Buckley’s discovery (1933) that some other species
of culicoides transmitted Mansonella ozzardi,
another non-pathogenic filaria, did no more alert
scientists to any potential problem. In animals, some
Onchocerca species are now known to be transmitted by Culicoides: O. gibsoni, a parasite of bovines
in Asia and Africa, and O. cervicalis and O. reticulata, parasites of horses.
Virus transmission in Africa
While the transmission of some protozoans has been
reported, the most important hazard of Culicoides
bites, from an epidemiological point of view, is the
transmission of viruses, even if no transmission to
man has yet been recorded. In the 1950s, the viruses of various animal encephalopathies were demonstrated to be transmitted by Culicoides in different
regions of the world. Last but not least, was the
description by R.M. Du Toit, in 1944, of the transmission of blue-tongue virus (now known to be an
Orbivirus belonging to the Reoviridae group) to
sheep, by certain species of Culicoides. The disease
was known in South Africa, where it was described
in 1880, although it existed there for centuries. It
was also found, at the beginning of the 20th century, in other parts of Africa, but did not spread
beyond the continent before the 1940s. Probably for
this reason, little attention was paid to the disease,
which is also known as “catarrhal fever of sheep”,
although it was responsible for heavy casualties in
sheep. The acute stage follows an incubation of
about a week, and includes severe respiratory symptoms together with fever, lethargy and anorexia.
Lung and pharynx oedema induce cyanosis, which
gives its name to the disease, even if oedema and
clinical signs are not limited to the tongue. The
whole upper respiratory tract is involved: oedema
and haemorrhagic lesions occur in the mouth as
well as in the nose. Depending on the virus
serotype, mortality in sheep varies from 20 to 80%.
Bovines, although serious as reservoirs of infection,
do not develop clinical disease when infected by
most serotypes.
The crossing of the Mediterranean sea
Little attention was devoted to blue tongue and even
less to its vectors while the disease gradually spread
over Africa. In the early 1950s, however, the disease
reached the Mediterranean sea coast, and attracted
attention from entomologists in north Africa institutions, such as J. Clastrier at the Institut Pasteur of
Algiers, who completed the identifications made by
Kieffer in 1958. This work, along with studies by de
Meillon in South Africa, resulted in the conclusion
that the main vector of the virus was Culicoides imicola, Kieffer 1913. M. Kremer subsequently published a new description which is now the basis of
the diagnosis for this vector. Although many other
species were suspected of being vectors, none was
then proved to be infectious, and attention focused
on C. imicola when, slowly, the virus spread northward, reaching parts of Portugal and Spain between
1956 and 1960. Greece was infected for the first
time in 1979, and again in 1998. Blue-tongue virus
includes 24 serotypes of which four are present in
the Mediterranean basin: serotype 2 mainly in the
west (Algeria, Sicily, Sardinia), and serotype 9 in the
east (Greece and the Balkans), but 4 and 16 are also
known to occur. Southern Italy is infected by
serotype 4, and Italian entomologists have also contributed to knowledge of the vector.
At the turn of the 21st century, the situation
appeared clear: occasional incursions of blue
tongue disease could happen where conditions permitted C. imicola to spread. It was possible to limit spread by vaccination, even if nothing proved
effective against the vector. Luckily, the species
involved is a hot climate insect which does not survive winter in the Europe, and its normal distribution did not appear to exceed the 40th parallel, limiting it to southern Spain, Italy and Greece. But
then the phenomenon of so called Global Warming
manifested itself. If climatic change was not admitted by all, some entomologists began to fear extended distribution of the insect vectors. In October
2000, C. imicola was found in Corsica, having
arrived from Sardinia, so confirming this risk and
the disease followed the insect, invading the whole
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251
Figure 1. Blue tongue fever invades Europe: situation during summer 2007.
island within few years. The Balearic islands were
similarly infected subsequently, demonstrating that
the vector can be carried long distances by wind. In
1995, P. Mellor published a map based on the temperature requirements of C. imicola, which detailed
the regions where the disease might already spread,
and predicted its reach should temperatures
increase. Mellor indicated a minimum year-round
temperature of 12.5°C as permitting the insect to
establish itself. With global warming, each additional degree Celsius would bring the vector 90
kilometres further north.
The threat was considered serious: the mediterranean regions, and Southern Europe generally, are
places of sheep farming. Blue tongue disease dissemination across the region would be a disaster. An
important program of surveillance and prevention
was set-up, based on entomological and veterinarian surveys. In France at least, it was very difficult to
find anyone knowing something of Culicoides.
There had for decades been a brilliant school of
entomology in Strasbourg, where J. Callot and M.
Kremer were renowned for their work on this subject. Unfortunately, both had retired before the issue
subject became topical: and the midges were not
attractive as a subject of research for their successors. None the less, the fortress was prepared to
resist the assault, looking southward to detect any
C. imicola invasion. Some were detected in 2005
near the Mediterranean sea coast, but no case of
blue tongue disease was detected. In the neighbouring countries as well, the disease remained limited
to the south and appeared to be under control.
A sudden and unexpected attack of blue tongue disease
in the North: an exotic virus carried by local vectors
Would the increase of temperatures in the region
really have permitted the settlement of C. imicola
and the spreading of the virus on the northern side
of the Mediterranean sea? Nobody will ever be able
to answer that question, as everything suddenly
changed in 2006: the European fortress was taken
from the rear! The disease entered Europe through
Belgium, and spread rapidly to the Netherlands, Germany and Northern France, although no C. imicola
were present in the region. The scenario is, however,
clear. The virus (probably imported from Kenya
along with living sheep), was a variant (serotype 8)
which happened to be transmitted not only by C.
imicola but also by widely distributed local European species, such as C. dewulfi and C. obsoletus
(Mehlhorn et al., 2007). Moreover, this variant was
more pathogenic for sheep, and also brought severe
clinical manifestations in bovines. Although drastic
measures of isolation and confinement were taken,
the disease spread rapidly during the summer of
2006. During that winter, populations of biting
midges were found indoors at Liege (Losson et al.,
2007) and a massive outbreak seemed possible during the following summer, in 2007.
This is exactly what happened. As the end of summer 2007 was mild, the vectors remained active
longer and the epizootic continued, and still continues to extend, overcoming any possibility of
blocking it. By mid-September, 23 departements in
the North of France were infected, and the epizootic continued to move swiftly southward during
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the autumn and early winter. In December 2007, 55
departments reported cases. The agricultural
authorities remained watchful, as reported in such
official websites as www.afssa.fr or www.agricul
ture.gouv.fr, but were entirely unable to arrest its
progress. In an attempt to prevent the introduction
of the disastrous variant 8 into their country, the
Italians forbade all imports of live cattle from
France. During the autumn also, cases were found
in England: it is well known that winds can carry
Culicoides long distances, much further than the
width of the Channel. The same progression
occurred in other Western European countries, from
the initial foci in Belgium and Netherlands the disease moved east towards Germany.
Nothing to do with global warming
There is nothing magic in this dramatic progression
of the disease. It was simply due to the unexpected
arrival of an infectious agent in a place where efficient potential vectors were already living, “flying
by thousands near of streams” as reported by the
Reverend Derham. As long as other variants of the
virus were involved, European species of Culicoides
could not spread them. Unfortunately, they were
susceptible to variant 8, which originates in East
Africa and had never encountered any European
species before arriving in Europe in 2006. In the
past, various other infectious agents found a similar opportunity, which allowed them to infiltrate
new areas of distribution. For example, when old
world visceral leishmaniasis arrived in South America, carried by Spanish and Portuguese dogs, the
parasite found a local sandfly which ensured its life
cycle and permitted it to invade a large part of the
continent. The same type of infective opportunity
occurred once again in South America with the
introduction of schistosomiasis mansoni carried by
African slaves.
On this occasion, the introduction of blue tongue
virus happened under the eyes of a community of
veterinarians specialised in infectious diseases, epidemiologists and virologists. Moreover, three centuries of entomological research had delivered the
key to understanding the background to the diffusion of this extraordinary epizootic, even if the distress of medical entomology had dramatically
reduced the number of people working in this particular field, and able to evaluate the risks of transmission. All the conditions for understanding the
parameters of the invasion were quickly identified,
but nothing could stop it. From an economic point
of view, the epizootic is a disaster in sheep breeding, but also for cattle. The vaccine against
serotype 2, which is produced in France and used
in Corsica, does not protect against the invader
serotype 8. A vaccine against this serotype however exists in South Africa. Now being prepared in
France, it is expected to be available in summer
2008.
It is certainly too early to draw conclusions from
this historical event. If it was a human disease
instead of an animal one, its importance would
already have exceeded any of the newly emergent
pathologies since AIDS. Fortunately, the species
barrier between ruminants and man has not (so
far?) been crossed in the blue tongue case, as it has
recently been by other viruses, such as AIDS or the
flu viruses. It is crucially important to recognize
that this spectacular zoonotic outbreak depends on
the same fundamentals as older epidemics. It
reminds us that, whether domestic animals or
humans, only experience can protect from epidemic threats, and that none of the knowledge gathered
in the past, even if it concerns studies on tiny
midges, can be considered useless in the encounter
with the world of pathogenic micro-organisms, particularly viruses, and emerging or re-emerging diseases.
Acknowledgements
The author wants to acknowledge the essential help received
from M. Kremer, who remains one of the best sources of information in the field of midges.
References
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Clastrier J (1958). Notes sur les Ceratopogonidés. IV. Ceratopogonidés d’Afrique Occidentale française. Arch Inst
Pasteur Algérie 36: 192-258.
de La Rocque S, Hendrikx P (2001). Impact du changement
climatique sur la santé. Exemple de la fièvre catarrhale du
mouton ou “blue tongue”. Changement climatique et maladies à vecteurs: Workshop in Nice, 18.11.2001, CR, p 62.
Derham W. Théologie physique ou démonstration de
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du Toit RM (1944). The transmission of blue-tongue and horsesickness by Culicoides. Onderstepoort J Vet Sci Anim Ind
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S, Gregory M. Les maladies émergentes consécutives au
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C. tropicalis Kieffer, sur des exemplaires déterminés par
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1972, Zoologie 44.
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ordinem naturalem in familias disposita, iconibus explicata.
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de Deken G, Deblauwe I., Fassotte C, Cors R, Defrance T,
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Mellor PS, Boorman JPT. (1995) The transmission and geographic spread of African horse sickness and blue tongue
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Mehlhorn H, Walldorf V, Klimpel S, Jahn B, Jaeger F, Eschweiler J, Hoffmazn B, Beer M (2007). First occurrence of Culicoides obsoletus transmitted bluetongue virus epidemic in
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Reconstructing an epistemological itinerary:
environmental theories of variation in Roubaud’s experiments on
Glossina flies and Anopheles, 1900-1938
A. Opinel
Centre de recherches historiques, Institut Pasteur, Paris, France.
Abstract. This paper addresses the theories and debates concerning the influence of environment on vectors and species variation. In particular, it focuses on theories about how climate and domesticated animals affected vectors that transmitted sleeping sickness and malaria. Emile Roubaud (1882-1962), a Pasteurian entomologist, worked on the adaptation and variation of Glossina fly races in order to elaborate
environmental interventions for sleeping sickness campaigns in Africa. He then developed the theory concerning Glossina flies’ biting preferences for livestock, and the implications of such preferences for human
protection against sleeping sickness transmission. Subsequently, he extended this theory about insect biting preferences to malaria in Europe. He thus used one disease model, the sleeping sickness complex,
and extended it to another, the malaria complex. He subsequently became interested into zoophilic races
of Anopheles maculipennis and advocated the hypothesis that the zoophilic Anophelines’ maxillary index
was a decisive feature in malaria transmission, for it could help preventing humans from the bite of the
Anopheles vector. The paper also analyzes how these theories were received and debated at the time of
their publication in scientific journals and proceedings.
Key words: Roubaud, medical entomology, environment, variation, Institut Pasteur.
The present research examines the Pasteurian entomologist Emile Roubaud’s early twentieth-century
research on and theories about insect adaptation
and variation. Emile Roubaud (1882-1962) (Fig. 1),
an university trained entomologist did not restrict
his work to observations of insects and parasites,
but designed theories and used the latter in designing experiments aimed at reproducing the variations
he had observed. He concluded from his observations that environment played a central role in variations among insects. Roubaud’s study of the environment and its influences on vector variation,
focused especially on climate (heat, humidity,
winds), but also on the living environment and biocenosis. We analyse the question of the contribution
of these parameters on the aetiology and distribution of diseases. Roubaud’s research and experiments on Glossina flies and Anopheles maculipennis will provide the models of our investigation
about his observations, experiments and theorization, that is to say his epistemological itinerary.
This paper is primarily based on his scientific papers
and correspondence. Roubaud engaged in an important correspondence, thus far unexamined by other
historians, with the Pasteurian Félix Mesnil (18681938)1 between 1909 and 1910, when he undertook
Correspondence: Annick Opinel, Centre de recherches historiques, Institut Pasteur, 28 rue du Dr Roux, F-75724 Paris
Cedex 15, France, Tel 33 (0)1 45 68 82 86, Fax 33 (0)1 45
68 81 84, e-mail: annick.opinel@pasteur.fr
1
Archives de l’Institut Pasteur (hereafter AIP), Mesnil
Papers. MES.04.
Figure 1. Emile Roubaud. Photothèque historique, Institut
Pasteur.
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A. Opinel - Roubaud’s environmental theories of variation
his second African mission. This correspondence,
the starting point of this essay, has provided the raw
material about his research on Glossina flies conducted during his African missions. The essay also
draws from a complementary literature, such as Bulletin de la Société de pathologie exotique, les
Annales de l’Institut Pasteur, entomological congress proceedings (the first in Brussels in 1910, the
1932 Paris congress and Amsterdam’s and Cairo’s
congresses in 1938), as well as materials from the
Institut Pasteur’s cours Roubaud d’entomologie
médicale.
Roubaud’s African missions: from the cycle of
trypanosomes in Glossina flies to the influence of
climatic conditions on sleeping sickness epidemics
Emile Roubaud was trained as a zoologist and began
to specialize in dipterology when he was trained at the
Museum national d’histoire naturelle, under the
supervision of the entomologist E. Bouvier 2. From
1905 to 1912, he was hired by Félix Mesnil at the
Institut Pasteur. His primary work, however, was conducted in Africa before he created his own laboratory, le Laboratoire d’entomologie médicale et de biologie parasitaire, in 1912. Roubaud’s doctoral thesis, La
Glossina palpalis, sa biologie, son rôle dans l’étiologie
des trypanosomes (1909), constitutes part of the
report he wrote after his first mission in Afrique équatoriale française (AEF, Congo) from 1906 to 1908.
Emile Roubaud was in charge of creating a Cours
d’entomologie médicale, the first of which should
have been given in 1912. But because of his missions
in Africa and the upheavals of World War I, the
course did not begin before 1922. At that time, it was
offered within the framework of the Cours de microbiologie. Roubaud’s course provided a detailed
description of each insect vector, and an analysis of
each infectious agent’s life cycle and the pathology it
provoked3. In 1926, the medical entomology section
of the microbiology course was transformed into a
Cours de protozoologie médicale. This pedagogical
structure persisted until Mesnil’s death in 1938.
Roubaud had thus genuinely been a dominant actor
of medical entomology at the Institut Pasteur.
After Emile Brumpt’s interrupted mission to study
sleeping sickness in Brazzaville in 1903 (Brumpt,
1934), the Ministry of Colonies became increasingly preoccupied by the extension of sleeping sickness
in Africa, which jeopardized the economic development as a large part of the population, already suffering from yellow fever, malaria, was as well victim
of sleeping sickness. Following the discovery of the
vectors of these diseases, several scientific and medical missions were organized between 1901 and
1914, most of them involving the Institut Pasteur.
Institut Pasteur’s involvement in these missions was
For his academic training, see Gachelin-Opinel, this issue.
AIP, Service d’entomologie médicale, Box SEM 1, Cours
Roubaud, fascicule 1, cours 1, p. 1.
2
3
the result of Roux’s willing. Emile Roux (18531933), a very influential man and director of the
Institut Pasteur from 1904, was a member of most
French hygiene and health commissions and he
advocated for Pasteur to become involved in these
missions. In some cases, state agencies requested
that Institut Pasteur participated in the missions.
Indeed, the Institut assumed scientific authority for
several of these missions 4. The sleeping sickness
missions organized in Congo in 1906-1908 and in
Senegal in 1909-1912 demonstrate the growing role
granted to entomology and research in controling
tropical diseases (see below). They also illustrate the
complexity of institutional networks involved in the
designing of the policy of control of endemic diseases. From a scientific viewpoint, they inspired
Emile Roubaud to begin thinking about the ways in
which specific environmental conditions could fundamentally shape epidemics, a question which preoccupied him until the end of his career.
The first mission was organized by the Paris-based
Société de géographie, which published the proceedings entitled La maladie du sommeil au Congo
français, with a preface by Emile Roux (MartinLeboeuf-Roubaud, 1909)5.
Sources for Roubaud’s specific research activities
and findings on this first mission are found in several publications. The report of the mission includes a
265-page chapter Biologie et trypanosome and
authored by Roubaud (a third of the entire book).
The chapter itself examines the combined entomological and protozoological results of his work in
Congo. Several other findings, not included in the
mission’s report, also brought together entomology
and parasitology and were published in journals
(Roubaud, 1935) or addressed in letters to his pasteurian chief Felix Mesnil. These available sources
indicate that Roubaud’s major findings during this
first mission concerned the biology of trypanosomes
inside the tsetse fly. As early as 1908, Roubaud had
already identified trypanosomes in the proboscis of
the fly. This discovery, along with his identification of
the importance of the vector’s saliva in facilitating
transmission, stand as the major contribution of his
research. Indeed, Roubaud performed numerous significant experiments that shed light on the parasite’s
movement and its complex development between fly’s
midgut, proboscis, and back to salivary gland.
Roubaud’s second mission (1909-12) to Africa (Senegal) with Médecin-major Georges Bouet was more
epidemiological in nature6. Roubaud and Bouet
4
Marchoux, Simond and Salimbeni’s mission to Rio 19031906 on yellow fever (Lowy, 2001) and Sergents Brothers’
long term mission in North Africa for malaria (Sergent, 1928;
Dedet, 2000).
5
The organisation of missions as well as the biological
instructions are detailed in A. Opinel (2008).
6
Roubaud and Bouet had worked together in Mesnil’s laboratory, and Bouet subsequently assumed responsibity for the
Dakar health service in 1913.
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sought to study the transmission of different endemic viruses by Glossina palpalis, morsitans and tachinoïdes, as well as these vectors’ respective roles in
transmitting various human and animal trypanosomiases. Indeed, the major goals of this second mission
were to follow the progress of both human and livestock trypanosomiasis epidemics, to study parasitic
diseases in animals (Spirilloses, Filarioses, Piroplasmoses), to investigate thoroughly tsetse fly biology,
and to find ways of destroying those insects.
Specifically, Roubaud and Bouet studied several
important medical entomological questions that had
been explicitely asked by the instructions they
received:
– Was endemic sleeping sickness transmissible by
the mean of Glossina flies?
– What were the conditions of endemicity of trypanosomiasis in tsetse-infested areas?
– What specific role did flies and domesticated or
wild animals play as hosts of viruses?
– How long did viruses incubate within Glossina
flies before these flies transmitted them?
– What role did secondary vectors, such as Stomoxas, horse flies, fleas and other biting insects,
play in spreading epidemics?7.
Roubaud and Bouet published their findings 8, in
which they identified nine species of Glossina flies
and four trypanosomes. Roubaud admitted in his
several letters that he was primarily interested in
studying the insect vector, since the parasite’s biological properties revealed only some physiological
adaptation of both the parasite and the insect to climate and nutrients9.
7
AIP, MES.04, A.S. de la mission Bouet-Roubaud, Dakar
le 13 août 1909, Med Inspecteur de service Callay to the
Gouverneur général de l’AOF. Here are the original instructions:
– La maladie du sommeil est-elle endémiquement transmissible par les Glossines déjà citées?
– Quelles sont les conditions d’endémicité des trypanosomiases dans les zones à glossines?
– Quel est le rôle des mouches et des animaux domestiques
ou sauvages dans la conservation des virus?
– Quel est le temps d’incubation pendant lequel les Glossines
gardent les virus avant de les transmettre?
– Rôle des agents secondaires de transmission: Stomoxes,
Taons, Moustiques, insectes piqueurs divers, comme
vecteurs des épidémies.
– Suivre la marche des épidémies de trypanosomiase humaine
dans les cases et des épidémies de troupeaux.
– Etudier les maladies parasitaires voisines des animaux
(Spirilloses, Filarioses, Piroplasmoses).
– Approfondir la biologie des Glossines et trouver le moyen
de détruire ces insectes.
8 Bulletin du Comité d’études historiques et scientifiques de
l’Afrique occidentale française, Gouvernement général de
l’AOF, 1920.
9 AIP, MES.04, Roubaud letters. Roubaud to Mesnil,
20/4/10: description of trypanosomes adhering to the exterior of Glossina fly salivary glands. Roubaud to Mesnil
20/3/10: discussion of geographical races of Glossina flies,
indistinguishable on a morphological basis, but differences
resulting from the insect’s adaptation to different tempera-
257
Given Roubaud’s avid interest in vectors and the
ways that environmental conditions shaped them, it
is no surprise that his third mission to Sénégal in
1913 (not discussed here but devoted to agriculture
problems due to termits) was strictly entomological
(Roubaud, 1935), and that his later work remained
primarily focused on insects.
Early experiments to control the virulence
of trypanosomes
Roubaud and his colleague Bouet arrived in AOF at
Thiès (Sénégal) in 1909, and subsequently traveled
to Agouagon (Dahomey, now Bénin) where most of
the correspondence to Mesnil has been sent from.
Roubaud pursued there the experiments initiated for
his Ph. D. thesis during his first African mission,
concerning the influence of saliva on Trypanosoma
development in Glossina flies. This constituted the
aim of his 1910 experiments such as transformation
of Tr. gambiense in digestive tube and, above all, the
influence of the salivary liquid in the transformation
of the shape of the trypanosome and the verification
of his earlier theory of the influence of climatic features on saliva properties.
The major challenge of parasitology, then and
now, centered on untangling the complex and interacting causes of illness, since parasites interact with
a wide range of human, animal, insect hosts, which
are also affected by broader climatic, seasonal, and
ecological influences. Roubaud, seeking to account
for these multiple causal factors, designed his experimental protocols so as to confirm the theory that he
elaborated through his observations, with controlled
experiments that allowed him to explore independently each climatic parameter. In order to conduct
these experiments, he used animals of different
species (goats, sheep, dogs, etc.), sound (healthy) or
infected, to test the ability of different Glossina flies
on different hosts (reciprocal species specificity). He
also sought to experiment on the various Glossina
vectors, either wild (“caught in nature”) infected
adults or laboratory-raised pupae or teneral adults.
Roubaud also found it necessary to seek out Glossina species of Glossina flies other than G. palpalis
such as G. morsitans. He further identified other
different Trypanosoma responsible for various animal and human diseases including “Trypanosoma
nagana, gambiense, surra, pecaudi”10. Roubaud’s
tures and hygrometry. A scientific project seeking to adapt the
Glossina fly to specific environments so that it could not
transmit Trypanosoma was also discussed.
10 AIP, MES.04, letter from Agouagon. “(…) une expérience en cours avec Schilling, Nagana, Gambiense, Surra,
Pecaudi” (letter dated 22 Feb 1910) or “Cette infection permanente de la trompe qui ne prend fin qu’avec la vie de la
mouche, je ne l’ai observé jusqu’à présent que pour Tr. cazalboui. Pour tous les virus avec les quels j’ai opéré: gambiense,
pecaudi, nagana Zoulouland et nagana Togo, surra M. (?), les
résultats ont été entièrement négatifs” (letter dated 20 March
1910).
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correspondence with Mesnil highlighted the crucial
importance of the chronology that he used to establish the incubation time of the trypanosome in the
body of the Glossina fly. Blood analyses on different
experimental animals established the diverse shapes
and stages of the parasite and their relative localisation11. Roubaud wanted to emphasize that the trypanosome was introduced into the vertebrate host
through the fly’s infected proboscis at the very
moment of the fly’s bite:
Au point de vue morphologique il y a un point intéressant à mettre en évidence (…). C’est la présence qui
paraît tout à fait constante de formes trypanosomes
normales et libres, aussi agiles dans le sang, localisées
exclusivement dans la lumière de l’hypopharynx. Je
l’avais observé au Congo dans le cas de ma mouche au
3° jour. Ici j’ai pu vérifier très nettement cette observation. Les trypanosomes qui sont en général en petit
nombre, voyagent librement dans la longueur du tube
excessivement fin de l’hypopharynx (…) Chose curieuse on ne trouve à côté d’eux aucune des formes Leptomonas, qui sont toujours localisées à l’extérieur de
l’hypopharynx et fixées aux parois du labre de préférence. Ainsi, les conditions de la salive à l’intérieur de
l’hypopharynx sont différentes de ce qu’elles sont dans
le reste de la trompe, et permettent la transformation
des formes fixées en formes trypanosomes. Ces trypanosomes bien entendu ne remontent pas vers les
11 African trypanosomes are sanguicolous flagellates with an
extra cellular development. The genera Leishmania and Trypanosoma include species that are pathogenic for human
beings. The genus Trypanosoma contains another human trypanosome, Tr. cruzi, responsible for Chagas disease. The two
trypanosoma subgenera that are pathogenic for human beings
are Tr. gambiense and Tr. rhodesiense (species group Tr. brucei). These sub-genera are distinguished on the basis of their
geographical distribution, their modes of transmission, and
the symptoms that they provoke. The trypanosomes Tr. brucei, Tr. congolense and Tr. vivax infect specifically animals.
Morphological differences are slight.
Trypanosomas develop following different sequences and
different stages in the vertebrate host’s blood and within its
digestive system (but not in all vectors). They assume different forms, according to their stage of development. Tr gambiense, for instance, takes on two forms within the Glossina fly:
the stumpy forms within the fly’s midgut, and the slender
form, within its salivary gland. [This dimorphism provoked
debates about whether trypanosomes underwent a sexual
cycle, a possibility that Roubaud himself examined in his thesis (Roubaud, 1909); he quoted Minchin, Gry and Tulloch’s
1906 work about Tr. gambiense transformation in the palpalis
digestive tube. In his thesis, he attributed changes following
ingestion to sexual differentiation into the male (slender) and
the female (stumpy) form. The three sub genera brucei, gambiense and rhodesiense can be compared on a morphological
level.
The cycle of the trypanosome in the tsetse fly lasts about 20
to 30 days. The fly first ingests slender forms during its blood
meal. Only the stumpy forms are able to carry on their evolution inside the vector; slender forms, in contrast, are responsible for multiplication and tissue invasion inside the mammal. The tsetse fly is not merely passive vector of the disease,
but rather facilitates the trypanosome’s evolution in its digestive system. Within the brucei species, parasite develops first
inside the midgut, then in the proboscis, and finally in the salivary glands of the tsetse.
glandes salivaires; on les trouve de préférence à
l’extrémité libre du tube hypopharyngien vers la sortie
de l’organe et par conséquent prêts à être inoculés.
J’imagine que ce sont surtout ces formes, chassés de
leur lieu d’élection par une poussée brusque de liquide
salivaire au moment de la piqûre qui sont les agents de
l’infection. Les formes Leptomonas sont si nettement
fixés aux parois de la trompe qu’elles ne peuvent être
guère déversés dans le sang au moment des piqûres.
Elles sont les agents du maintien de l’infection dans les
trompes: les trypanosomes sont des formes propagatrices à l’extérieur 12.
These lines constitute one of Roubaud’s most important contributions to the epidemiology and transmission of the disease: he contended that scientists
needed to identify the specific location of the infectious agent within the vector, and he hypothesized
that this site was the fly’s proboscis. Most important, he linked the parasite’s evolution with the
influence of saliva in the proboscis.
Roubaud thus defined the complex causation, the
intertwining of cause and effect. Because of the various local climatic conditions (dryness, humidity,
and heat), the Glossina fly saliva is modified and
acquires different properties, which in turn affect
the proliferation of Trypanosoma inside the vector
and ultimately, local differences in infectivity. It was
this complex causation that Roubaud asserted he
has proven:
Je vous disais dans une précédente lettre que j’étais
presque sûr d’arriver à faire perdre leur pourvoir infectant à des mouches au cazalboui en les soumettant à
des conditions de température ou d’humidité différentes de celle qui règnent ici. Cette idée a été parfaitement vérifiée par l’expérience (…) Tr. cazalboui est
certainement à cette latitude le virus de choix pour
faire quelques expériences. C’est un excellent réactif
pour apprécier la sensibilité physiologique des glossines à l’influence de la sécheresse. Je recommencerai
ces expériences à plus grande échelle13.
In one experiment, Roubaud confined wild tsetse
flies to a container of air slightly dried with potash.
The flies subsequently mated, laid eggs, and were
then fed on a young goat infected with Tr cazalboui.
Roubaud found that nine days after their last infected blood meal, the flies confined to dry air contained no trypanosomes at all. In contrast, flies confined to normal (humid) laboratory conditions
“would have been infected in proportion of 7 to 8
out of 10”. Roubaud thus concluded that the degree
of humidity fundamentally affected the flies’ transmission capacity, by altering the properties of trypanosomes, either in number or in infectivity.
Roubaud asserted in his 1909 doctoral thesis: “les
phénomènes de multiplication qui se passent dans
l’intestin des mouches, au labo, dans des conditions
ordinaires, sont de simples phénomènes de culture”
12 AIP, MES.04, letter from Roubaud to Mesnil, Agouagon,
20 March 1910.
20 April 1910.
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259
Figure 2. Afrique occidentale française, Mission Bouet-Roubaud, 1906-1910, Trypanosmiasis and tsetse flies distribution map. In:
Bulletin du Comité d’études historiques et scientifiques de l’Afrique occidentale française, Gouvernement général de l’AOF, 1920.
(Roubaud, 1909). In 1910, however, he revisited
this claim:
“Le retour aux formes trypanosomes indique bien un
cycle évolutif, mais qui est complet, comme je l’ai
indiqué autrefois, dès les premières heures, pour certains virus. Le terme de culture appliqué aux phénomènes qui se jouent dans la trompe est véritablement
impropre. C’est une évolution biologique qu’il faut
dire (…) Je crois en effet maintenant pouvoir donner
des ‘infections totales’ une interprétation inverse de
celle que je leur ai donnée naguère. Ce ne sont pas les
formes de l’intestin qui doivent remonter vers la trompe, mais bien plutôt les formes salivaires qui, entraînées dans l’intestin par déglutition de la salive, y propagent une culture durable entretenue continuellement par la salive infectée et qui n’a rien de nécessaire pour la transmission”14.
This passage reflects both Roubaud’s frank willingness to re-think earlier claims. He acted as an experimental entomologist, anticipating an experimental
approach concerning environment factors.
The following apparently anecdotic episode shows
the importance he granted to the observed fact and
to the logical consequences which can be drawn
from it. He openly admitted his confusion or his
hesitation, expressing surprise, when two G. tachinoides were born from pupae in a laboratory cage
14
AIP, MES.04, letter from Roubaud to Mesnil, Agouagon, 20
March 1910.
ostensibly containing G. palpalis. He wondered if
perhaps he had accidentally captured a few tachinoïdes, or if it were a brutal variation of palpalis15.
The possibility of a contamination was high. However, in the context of French biology of the time,
Roubaud seriously considered a brutal variation of
species as a possibility (Gachelin/Opinel, this issue).
It is not difficult to see from the above reflections
that Roubaud was thus predisposed to explore the
influence of environmental difference on Glossina
flies. He developed a map with Bouet (Fig. 2) to
illustrate the geographic distribution of Glossina
flies. His correspondence reveals his observations
about vegetation, heat, shadow or sun exposures
and their consequences for tsetse flies’ transmission
capacity leading to the notion of geographical races
which Roubaud often referred to:
“Je crois que l’on peut affirmer que les parasites sont
électivement adaptés à certaines races géographiques
des mouches, différant les unes des autres par leur
faculté de résistance à l’humidité ou à la sécheresse
ou mieux par les modifications que ces facteurs
apportent à leur milieu salivaire”16.
22 May 1910. “S’agit-il d’une variation brusque de palpalis?”.
16 AIP, MES.04, letter from Roubaud to Mesnil, Agouagon, 20
March 1910.
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These experimental insights enabled Roubaud to
develop recommendations to eradicate these vectors.
He advocated, for instance, clearing palm groves harboring Glossina flies rested; he underscored the role
of the harmattan, the hot dry season wind, in affecting Glossina flies’ biology. The map in Fig. 2 displays
the zones in which tsetse flies live and breed permanently (gîtes permanents), the routes of their dispersion, and their influence on transhumant pastoralist
economies. Roubaud thus sought to understand how
geographical zones and seasonal variations affected
the biology of trypanosomiasis vectors.
In a letter to Mesnil in 1910, Roubaud suggested
that tsetse fly races were undistinguishable on a
morphological basis, but rather their differences
resulted from their adaptation to different temperatures and hygrometry17.
In such defining races, Roubaud raised critical
questions about the heterogeneity of insect populations. The taxonomy of Glossina (genus) had
already been described in 1830, and subsequently
the species G. morsitans and palpalis were identified, distinguished on the basis of their morphological features (Cambefort, this issue). Roubaud introduces here the notion of biological races, identical
in morphology within each species but different by
biological criteria that can only be appreciated in a
given environment. Roubaud thus not only deepened entomological understanding of the biology of
the Glossina fly, but he also identified new features
that served to refine its existing taxonomy.
Roubaud continued to refine his theories concerning
environmental influences on vector transmission
capacity, articulating a concept of “conditions of
receptivity” (“conditions de réceptivité”). The “conditions de réceptivité” of parasites referred to the
infectivity of the vector’s salivary glands. On the
basis of his experiments in Moyen-Congo (Brazzaville), Moyen-Dahomey (Agouagon), Haute
Casamance (Kolda), Roubaud argued that variations
in receptivity resulted from geographical influences.
Also, he contended that the virus specifically adapted to Glossina species (longipalpis and palpalis: Tr.
cazalboui; Tr. dimorphon to longipalpis and tachinoides, etc.). Roubaud therefore established that the
“elective adaptation of certain viruses to certain
species of Glossina flies in some defined regions was
the result of modification of flies ‘receptivity’, following the biogeographical influences to which they
have been subjected to” (Roubaud, 1913, p. 30). He
stressed his argument by citing British, Belgian, German colleagues who have been studying geographical distribution of insect and diseases since he
first set up this notion in 1910 (Roubaud, 1913)18.
17
MES.04, letter from Roubaud to Mesnil, Agouagon, 20
April 1910.
18
Kleine in Eastern Africa, Bruce and his collaborators in
Uganda, Kinghorn and Yorke in Rhodesia, Kleine and Fisher
on Victoria lake banks and Taute, Lake Tanganyika.
He thus concluded that “the receptivity of a determined specie of Glossina fly for a given virus is not
a uniform property in the whole living area of the
species” (Roubaud, 1913, p. 32). Races of Glossina
flies displayed regional distinctions and different
degrees of receptivity. Roubaud also emphasized
that climatic factors did not directly influence the
virus’s evolution inside the Glossina fly; rather, biogeographical factors modified specific physiological
conditions within the salivary gland, which in turn
played upon the virus. Roubaud’s arguments about
zones of receptivity had important implications for
fly control measures, particularly that of brush
clearing. These receptivity zones, he argued, needed
to be taken into account in order to render such
measures more effective 19.
Great mammals as an environment for
human and vectors: livestock and
human protection against Glossina flies
Roubaud’s concept of environment as a significant
set of influences on sleeping sickness was not limited to vegetation, soil, and climatic conditions. He
also included animals, particularly those sensitive to
the biting of Glossina flies. Over time, Roubaud
articulated a theory concerning how animals could
attract vectors away from humans, and he called
this phenomenon “la méthode trophique” or “prophylaxie trophique”. It was both a theory and a
means of prophylaxis that he advocated throughout
his career 20.
Roubaud’s theory of trophic prophylaxis was
based on a simple observation: higher densities of
Glossina flies meant lower infection rates. He thus
assessed that the high rate of the disease was
inversely proportional to the abundance of tsetse
flies. Hence, the high rate of trypanosomiasis was
inversely proportional to the abundance of tsetse
flies. In Ubangui Chari, Bas Congo and Haute Sangha, Roubaud had found very high rates of trypanosomiasis, even though Glossina flies lived great
distances from human settlements. He explained
this phenomenon by contending that large mammals
normally provided a feeding source for vectors, and
that human beings served as only secondary hosts.
Where livestock such as cattle, horses, and donkeys
were abundant, flies were numerous, and the risk of
infection was reduced for human beings, because
livestock served as provided blood meal sources for
flies. But in central Africa, few livestock existed to
draw Glossina flies away from human beings, and
thus although Glossina were relatively scarce,
19
The same arguments can be found in a 1919 paper written by a Belgian entomologist, who described the geo-botanical conditions of the “gîtes à pupes” of Glossina palpalis, fusca, brevipalis, pallipides and morsitans. (Schwetz, 1919).
20 Roubaud had already argued that animals served as protection for human beings against malaria in his doctoral thesis; Grassi subsequently confirmed his theory.
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human populations constituted the primary source
of blood meals. Roubaud, however, was well aware
of the double-edged nature of this theory: livestock,
though they could draw Glossina flies away from
human beings, could also keep the flies alive. Efforts
to use livestock as “trophic prophylaxis” could thus
sustain infective Glossina feeding sources. He
sought in vain to disprove this counter-argument,
but ultimately he concluded that a balance had to be
struck between the “advantages and disadvantages”
(Roubaud, 1920a, p. 309) of the method in each
case!
Roubaud pushed this hypothesis even further.
Because big game constituted a natural reservoir of
diverse trypanosomes, they jeopardized domestic
cattle breeding. Hence large game populations
were antithetical to colonial interests. He consequently suggested a progressive replacement of
large game with domestic livestock, which could
then attract infective Glossina flies. Roubaud also
envisioned distributing domesticated livestock races
that were resistant to sleeping sickness throughout
Africa.
The biology and the anatomy of the Glossina flies
were for Roubaud the key to understanding the
transformation and the evolution of the parasite,
and in that respect, his concern about environmental parameters (vegetation and climate) was the other part of the explanation.
He subsequently sought to apply these findings to
malaria, going from the sleeping-sickness complex
to the malaria complex21.
Anopheles zootropic differentiation and
the maxillary index
Following his research on human African trypanosomiasis and tsetse flies, Roubaud devoted
much of his research to the malaria vector Anopheles, although he began to explore other insect vectors, as well as innocuous insects, such as Culex and
Locusta. His research following the first World War
was almost exclusively laboratory work, conducted
in Paris.
Beginning in the 1920s, Roubaud, as well as Marchoux and Sergent, became interested in the notion
of anophelism without malaria. This phenomenon
had been previously described by the Italian school
of malariology. Roubaud, drawing from his earlier
research and theorization concerning “trophic prophylaxis”, argued that the Anopheles maculipennis
maxillary frame (armature) could serve as an indi21 I refer here to Max Sorre’s work, in which he defined his
“pathogenic complexes (complexes pathogènes) (…) which
aggregates in the natural living environment (milieu) and
which possess their ‘global ecology’” (…) their synecology.
For Sorre, both complexes had the same general structure, “a
narrow adaptation of the parasite to the man and to the vector”, and he emphasized “the solidarity of those three terms”
(Sorre, 1947, p. 301)
261
cator of the degree of the zoophily among Anopheles races 22.
Roubaud developed and refined this theory in several published papers 23, emphasizing how confined
livestock could serve as a biological screen for human
beings against malaria. He asserted that malaria was
reduced when livestock screened human beings from
infective mosquitoes, and that Anopheline mosquitoes themselves altered in response to the presence of
livestock. Indeed, Roubaud, by analyzing the evolution of Anopheles zootropism, described a physiological evolution marked by “appreciable morphological
particularities on the different races” (Roubaud,
1921). He became interested in the study of maxilles
(see Danish researcher Wesenberg-Lund’s work 24)
establishing the axiom that the greater a mosquito’s
adaptation to livestock, the stronger its maxillary saw
would be. The average number of teeth, what he
called the maxillary index (the total number of teeth
divided by two), would be greater in a zootropic race
(that is, fauna with stronger zootropism), which
needed to perforate tough animal skins to obtain a
blood meal (Fig. 3). Roubaud contended that a maxillary index equal to or greater than 15 was characteristic of a zoophilic race, although he based his
study on a very small sample of anophelines. Thus, a
high maxillary index of Anopheles would induce low
malaria infection rates within human populations and
therefore had a predictive value.
Most interesting are the responses Roubaud’s theory elicited within the scientific community. For
example, as in 1922, the Sergent brothers, along
with Parot and Foley (Sergent et al., 1922),
Roubaud’s colleagues from Institut Pasteur d’Algérie,
published a paper in the same journal as did
Langeron (Langeron, 1922) from the Faculté de
médecine de Paris, laboratory of parasitology. Both
papers were very critical about the maxillary index
theory. Sergent et al. had conducted a survey in
Algeria on 1222 female A. maculipennis to determine whether they could confirm the Roubaud’s
theory. Their conclusions were quite clear: maxillary
teeth general average in the plaine of Bône was
14,4; the relative proportion of Anopheles with over
14 teeth was 46,4 percent. Following Roubaud’s
theory, these results should have indicated that the
plaine of Bône was a zone without malaria. But this
was far from the case, and the Sergents concluded
that “the theory (could) not be confirmed in Algeria” (Sergent et al., 1922).
Langeron found Roubaud’s theory an audacious
and seductive one, but asserted that it could not be
tested. Langeron offered a starkly different envi-
For the context of this theory, see Fantini (1994).
Annales Institut Pasteur, 1920; BSPEx, 1921, 1922,
1925; Congrès du paludisme, Alger, 1930; Congrès
d’entomologie de Paris, 1932; Amsterdam, 1938.
24 Wesenberg-Lund, Mémoires de l’Académie royale des
Sciences et Lettres du Danemark, 8e série, t. VII, n. 1, 19201921, quoted in Roubaud, 1921.
22
23
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Figure 3. Numerical variations in denticulations of the maxillary saw in Anopheles maculipennis (from top to bottom: 13
teeth, 15, 18). In: Roubaud E (1921), La différentiation des races zootropiques d’Anophèles et la régression spontanée du
paludisme. BSPEx 14: 582.
ronmental hypothesis: if Anopheles were found in
high concentrations within domesticated animals’
stables, these concentrations were due to the darkness and relative humidity that they required. Mosquitoes preferred these stables only because human
houses were taller, larger and better ventilated.
Langeron’s best argument against this theory was a
methodological one: he said that Wesenberg-Lund’s
and Roubaud’s hypotheses about maxillary denticulation and “other numerical data” remained interesting but they relied on far too small a sample
(from 4 to 25) to be convincing. In contrast to
Roubaud, then, Langeron advocated malaria control through more generally accepted methods of
the day, including distributing quinine, promoting
good human health (Langeron, 1922; Marchoux,
1921) and draining ponds and swamps. Roubaud
replied to these critics in an “acrobatic” way, admitting that his sample size was small, but insisting on
the value of his theory. Indeed, he contended that
drainage and the reduction of stagnant waters actually favoured Anopheles reproduction, which developed in permanent breeding places. They did not
disperse and breed next to hosts: “From wild, fauna (mosquitoes) becomes domestic” (Roubaud,
1922). Females thus altered their feeding habits,
developing a stable (zoophilic) preference for
domesticated animals. Wild Anopheles mosquitoes,
in contrast, unaccustomed to differentiating
between human and animal sources of blood meals,
would feed off of any source of blood that it
encountered. Thus Roubaud developed a rather tortured argument to defend his theory, explaining that
malaria reductions in drained areas came from the
increased presence of cattle, not from the suppression of mosquito breeding sites.
A definitive blow to Roubaud’s theory came in
1930, at Algiers Second Malaria Congress. There
Frédéric Trensz, a Pasteurian colleague from Institut
Pasteur d’Alger, published a study (Trensz, 1930)
that he had carried out with Edmond Sergent from
1928 to 1929, in Algeria, France, Italy and Spain.
The study tested whether Roubaud’s theory maxillary index could predict zoophilic behaviours among
mosquitoes, and thus be used to identify zoophilic
mosquito races. In his lengthy paper, Trensz, quoting in his bibliography about ten papers published
on this matter between 1919 and 1928, objected to
Roubaud’s other theories about maxillary index, that
is modification of maxillary frame under zoophilic
influence. The theory was considered weak. In Holland, for instance, malaria persisted, despite the
abundance of domesticated animals and the preference of Anopheles mosquitoes for these mammals.
Trensz first summarized Roubaud’s theory which
now includes the notion of “vital competition and
Darwinian ideas of natural selection”. Thus, as
Trensz understood it:
– When the cattle hosts are numerically insufficient,
Anopheles continues to bite human beings, since
it cannot be fed sufficient bloodmeals from livestock. The best armed individuals will be fed,
inducing an increase in the maxillary index.
– Fauna (mosquitoes) under intense competition
have few fed (gorgées) females and a high maxillary index.
– High indexed individuals bite more easily than
weaker ones.
– Malaria manifestations resume in populations of
highly indexed Anopheles. The existence of vital
competition is revealed by index over 15 (up to 14
and superior to 15 transmit malaria).
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– There is a difference of Anopheles maxillary
index, depending on host teguments: it could exist
several zoophilic Anopheles’ dental types because
of animals different teguments thickness and thus
“a selection leading to the creation of differently
armed races beside livestock of different species
(cattle, horses, pigs) in a same village”. Animal
skin thickness constituted a selective pressure on
mosquitoes, and there would develop in the same
village different mosquito races specialized to feed
off of different kinds of livestock.
Having defined Roubaud’s hypothesis, Trensz and
Sergent detailed their survey in the congress proceedings with elaborate diagrams and precise explanations, and concluded that in Algeria, Anopheline
mosquitoes possessed the same buccal frame,
regardless of the conditions that would favor or disfavor the development of zoophily. In Italy
(Bologna, Ferrara, Ravenna), they found that among
local mosquito with several dental types, no correlation existed between the presence of livestock and
the absence or presence of malaria. They reached
the same conclusions in France. The maxillary index
study of a great number of A. maculipennis (4244)
from Algeria, France and Italy did not bring any element in favour of a correlation with zootropism.
According to Trensz, the sample should have been
nearer to 10,000 individuals to be relevant. Hence
the survey did not confirm Roubaud’s theory.
What is fascinating about the discussion following
Trensz’s presentation of his paper at the Algiers congress is that despite the overwhelming evidence that
Trensz and Sergent had presented on the value of
maxillary index and the existence of inheritable food
habit, the audience remained in favor of the protective role of livestock.
The same 1930 congress saw the presentation of
another paper that provided a strikingly different
approach to environmental influences on mosquitoes
and malaria transmission, de Bruck, Schoute and
Swellengrebel presented their new research on
Anopheles maculipennis races in Holland and their
relation to anophelism without malaria (de Bruck et
al., 1930). Examining wintering and non-wintering
mosquito races, their wing sizes, and maxillary
indices, the Dutch researchers sought to determine
whether different environments (brackish water for
non-wintering, fresh water for wintering) provoked
the biological and morphological differentiation of
the two mosquito races. The authors’ argument countered Roubaud’s statement that races are mere modifications provoked by the environment (milieu), but
they asserted that the biological difference between
the two races persists when they are bred in the same
milieu. The ideal of the antimalaria fight was to dispose of the anophelian race by changing the environment (milieu), so as to favour the innocuous race 25.
25 “Car les changements du milieu ambiant, qui doivent
influencer, dans un sens opposé, les deux races d’une seule
263
Not all researchers at this meeting were equally
enamored with Roubaud’s theory. Alberto Missiroli
and Lewis W. Hackett thought that the theory of
“zootrophic deviation” could not be accepted, and
Erich Martini expressed doubts about differentiation
of zoophilic races. These criticisms and doubts
sparked the three researchers to undertake three
years of field research in the whole Europe to identify Anopheles maculipennis races, resulting at the
end of the discovery in 1935 of the maculipennis
complex (Fantini, 1994), which ruins the notion of
environment determined race (see below).
In the meantime, despite these criticisms at Algiers,
Roubaud staunchly defended his theory. In 1932, at
the fifth international congress of Entomology in
Paris, Roubaud gave a paper entitled Experimental
outlines on trophic and biological races of Anopheles maculipennis (Roubaud, 1932). In response to
the discussions in Algiers, he stated that he was
pleased that his thesis of Anophelian zoophilic deviation had sparked further studies about mosquito
differentiation. He referred to Martini, Missiroli,
and Hackett‘s research, which confirmed the existence of Italian races of Anopheles with different
zoophilic and anthropophilic behaviours.
Swellengrevel and van Thiel, too, had developed
theories based on the existence of two anopheline
types, a long-winged type that did not transmit
malaria, and a short-winged (atroparvus) that did 26.
Roubaud’s position remained unchanged concerning
the minor place he attributed to morphological features: “there are races of Anopheles maculipennis
differentiated and selected in a zoophilic sense and
indifferentiated races”. The races preferring humans
are paucidented, but, he said, the morphological
point of view must remain secondary. “Races of
Anopheles, zoophilic or not”, he argued, “must be
judged, before all, biologically” (Roubaud, 1932).
According to his experiments conducted in the
Institut Pasteur insectarium 27 on different strains of
Anopheles maculipennis. Roubaud stated that
trophic characters (i.e. breeding behaviours), similar
to maxillary frame general characteristics, are race
characteristics that were transmitted through
“heredity” 28 that were acquired by long exposure to
a given target (Gachelin-Opinel, this issue).
espèce d’Anopheles, seront évidemment plus faciles à provoquer que ceux visant deux espèces différentes” (de Bruck et
al., 1930, p. 299).
26 The study of short-winged mosquitoes constituted the
very first steps towards the identification of the maculipennis
complex.
27 In 1930, Roubaud built an insectarium at the Institut Pasteur in Paris following his mission to Tunisia with J. Colas-Belcour. This mission was comissionned by the section of
Hygiene of the Society of Nations (1927) to set up his “methods of experimental education” on different insects such as
Culex pipiens or Anopheles maculipennis. (Roubaud, 1935).
28 Missiroli assisted by sending several hundred of A. maculipennis from Toscana and Marshes, as well as from Spain,
Holland, Algeria, London, Vienna, and Normandy.
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Roubaud drew several conclusions from his experiments on trophic attraction:
– There exists trophic races of A. maculipennis
– There exists two large groups of Anopheles:
zoophilic group and anthropophilic group.
– There is a strong correlation between the maxillary index data and the experimental data which
permit the distinction between a population with
differentiated zoophilism and that with undifferentiated zoophilism.
– Paucidented strains of population with undifferentiated zoophilism are more easily attracted to
human beings and are barely engorged on human
blood meals than Anopheline races with with
higher maxillary indices.
– Similar to maxillary characteristics, trophic characteristics of mosquito population are race characteristics, hereditarily transmitted.
– There is a trophic memory (when Anopheles is
used to bite cattle, for example, it remembers it in
some way and comes back preferentially to cattle
for some time).
He sought to describe several other biological characteristics of the different mosquito races, including
reproductive behaviours, aggressivity of zoophilic
populations, higher ability of certain strains to
become fat 29 when grown on sweet food. In his
1932 paper, Roubaud distinguished between heterodyname (wintertime diapause, or dormancy) and
homodyname strains (reactivated by heat in wintertime), and he highlighted the ability of the hibernating females to undergo reactivation under the
effects of heat. The conditions of the ovulation diapause (asthenobienose) are thus not the same for
every population of Anopheles. He distinguished as
well stenogame race (στενός: narrow), which was
adapted to “domestic sedentarity”, and eurygame
race (εΰ̣ρυς, vast), adapted to open air population.
He further emphasized the ability of the former to
mate in limited spaces, including houses and of the
latter to mate outside (because they need a nuptial
parade before mating).
Roubaud concluded on the coexistence of heterodynamis with stenogamy and homodynamis with
eurygamy, as well as the influences of these combinations of characters on females’ hemophagous
activity, and in turn on malaria transmission:
Roubaud inferred that mosquitoes’ adaptation to
cattle appeared to be due to stenogame and heterodyname females, although he contended that the
great Dutch Anopheles race clearly belonged to the
zoophilic eurygame and homodyname type.
The scientific community, particularly Hackett and
Missiroli after their three years mission, resolved the
enigma of anophelism without malaria around
1935, when different Anopheles maculipennis subspecies endowed with different biological properties
each selected independently. The balance between
populations of different species explained the different properties of different field populations (Fantini, 1994). At least until 1935, Roubaud clashed
with other scientists, defending his theory concerning the environment’s promotion of species adaptation, the establishment of races, and the predictive
value of the maxillary index. The identification of
the Anopheles maculipennis complex did not stop
discussions about the existence of biological races,
now shifted towards the origin of these races, the
mechanisms at work, and the significance of the
maxillary index, mention of which can be still found
in the 1950s. The period between 1935 and World
War II witnessed the progressive decline in the
explanatory power of Roubaud’s theory concerning
biological races that had resulted from environmental influences. Roubaud’s lectures given after 193830
in his Cours d’entomologie médicale appear to synthesize his work, and mark the end of Roubaud’s
usage of “adaptation” as a paradigm for evolutionary processes and the generation of races. These lectures were more a description of the present features of insects than a practical course on medical
entomology 31. The underlying philosophy of the lectures appeared less combative than in his earlier
publications. Adaptation had not retained the evolutionary meaning that it had in the early 1930s. By
the late 1930s, it served more as an observation
than claim about a process (see Gachelin-Opinel,
this issue).
However Roubaud’s theories about variation within insect populations are now perceived, it is important to emphasize the impact of these theories on
scientific debates at the time. Roubaud’s theories
may have been wrong, widely criticized, or weakly
supported, but they nevertheless constituted an
important and original contribution to debates on
sleeping-sickness and malaria: they refined the taxonomy of Glossina flies and Anopheles by introducing biological, behavioural and environmental features to the description of species.
The question of the zootrophic deviation in the
fight against malaria was debated in scientific journals and in congresses for nearly 15 years. It can be
concluded that despite his great epistemological openess, Roubaud remained convinced about zootrophic
deviation, a theory first developed through his work
In this context, to grow fat (engraissement) means faster
development of fat bodies. Roubaud underscore that this biological specificity had also been emphasized by Swellengrebel
et al. as one of the significant biological characteristics differentiating the Dutch longwinged A. maculipennis and the
short-winged Anopheles.
30
AIP, box SEM.1, Recherche scientifique coloniale. Laboratoire d’entomologie médicale et zoologie tropicale, Institut
Pasteur. The exact date of the document is unknown; the most
recent reference given in the manuscript is dated 1937.
31
The lectures are based on Roubaud’s previously published
papers and five of them were devoted to the “adaptation
hémophage” and to feeding behavior.
29
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on tsetse flies prior to World War I. His synthetic
1920 paper “The trophic method in the fight against
insects and infections they carry” (Roubaud, 1920b)
extended his theory of animals as protective screens
for human beings against a range of parasitic diseases, including malaria, diverse trypanosomiases
(American and African), yellow fever, tick borne
fever, and afflictions transmitted by blood sucking
insects. He continued to develop and to provide evidentiary support for his theory until World War II.
In so doing, Roubaud articulated an almost utopian
vision of a long-term “trophic education of insect
parasites”, a “new biological ideal”.
Acknowledgements
I thank Tamara Giles-Vernick and Gabriel Gachelin for their critical reading of this essay.
References
Brumpt E (1934). Titres et travaux scientifiques. Paris: Masson
et Cie.
Dedet J-P (2000). Les Instituts Pasteur d’outre-mer. Cent vingt
ans de microbiologie française dans le monde. Paris:
L’Harmattan.
de Bruck A, Schoute E, Swellengrebel N-H (1930). Nouvelles
recherches sur les races d’Anopheles maculipennis au
Pays-Bas. Compte-rendu du Deuxième Congrès international du paludisme, Alger, 19-21 mai 1930, Institut Pasteur,
Secrétariat général du Congrès I: 293-300.
Fantini B (1994). Anophelism without malaria: an ecological
and epidemiological puzzle. Parassitologia 36: 83-106.
Gachelin G, Opinel A (2008). Theories of genetics and evolution and the development of medical entomology in France
(1900-1939). This issue.
Langeron M (1922). Sur l’anophélisme et le paludisme en France. Bulletin de la Société de pathologie exotique (hereafter
BSPEx) 15: 20-36.
Lowy I (2001). Virus, moustique et modernité. La fièvre jaune
au Brésil, entre science et politique. Paris: Archives d’histoire
contemporaine.
Marchoux E. Influence du bien-être sur la régression du paludisme. BSPEx 14: 455-459.
Martin-Leboeuf-Roubaud G (1909). La maladie du sommeil au
Congo français. Société de géographie. Masson, Paris.
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Opinel A (2008). The emergence of French medical entomology: the respective influence of universities, Institut Pasteur
and army physicians (ca 1890 to 1938). Med Hist 52(3): 387405.
Roubaud E (1913). Relations bio-géographiques des Glossines
et des Trypanosomes. BSPEx 6: 28-34.
Roubaud E (1920a). Les conditions de nutrition des Anophèles
en France (Anopheles maculipennis) et le rôle du bétail dans
la prophylaxie du paludisme. Annales de l’Institut Pasteur 34:
181-228.
Roubaud E (1920b). La méthode trophique dans la lutte contre
les insectes et les infections qu’ils transmettent. Revue générale des sciences pures et appliquées 31: 301-313.
Roubaud E (1921). La différentiation des races zootropiques
d’Anophèles et la régression spontanée du paludisme.
BSPEx 14: 577-595.
Roubaud E (1922). À propos des races zoophiles d’Anophèles.
BSPEx 15: 36-39.
Roubaud E (1925). Les raisons de l’absence en Europe septentrionale de l’endémie palustre estivo-automnale (Plasmodium praecox). BSPEx 18: 279-287.
Roubaud E (1930). Quelques remarques à propos de
l’interprétation théorique des index maxillaires. BSPEx 13:
47-53.
Roubaud E (1932). Aperçu expérimentaux sur les races trophiques et biologiques de l’Anopheles maculipennis. 5e
section: entomologie médicale et vétérinaire, V congrès
international d’entomologie, Paris, 18-24 juillet 1932, 715733.
Roubaud E (1935). Titres et travaux scientifiques. Masson,
Paris.
Roubaud E (1938). Les équilibres biologiques dans l’étiologie
du paludisme. Actes du congrès d’Amsterdam, 120-139.
Schwetz J (1919). L’identité des conditions géo-botaniques
des gîtes à pupes de la Gl palpalis. BSPEx 12: 234-239.
Sergent Edm and Et et al. (1922). L’armature maxillaire des
Anopheles maculipennis en pays paludéen. BSPEx 15: 2930.
Sergent Edm et al. (1928). Vingt-cinq années d’études et de
prophylaxie du paludisme en Algérie. Archives de l’Institut
Pasteur d’Algérie VI, 2-3.
Sorre M (1947). Les fondements de la géographie humaine.
Armand Colin. Paris.
Trensz F (1930). L’index maxillaire d’Anopheles maculipennis et
la théorie du zootropisme anophélien. Compte-rendu du
Deuxième Congrès international du paludisme, Alger, 19-21
mai 1930, Institut Pasteur, Secrétariat général du Congrès I:
155-225.
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Theories of genetics and evolution and the development of
Medical Entomology in France (1900-1939)
G. Gachelin 1, A. Opinel 2
Rehseis, UMR, 7596 CNRS - Université Denis Diderot, Paris, France;
teur, Paris, France.
1
2
Centre de recherches historiques, Institut Pas-
Abstract. The development of entomology and medical entomology in France is discussed in the context
of the prevalence of Lamarckian ideas concerning heredity and evolution. Lamarckian ideas have greatly influenced research carried out at the Institut Pasteur by Emile Roubaud and more generally in Felix
Mesnil’s laboratory, as well as research in general entomology at the Museum national d’histoire naturelle.
By contrast, it did not influence research and teaching at the Faculté de médecine of Paris or that of physicians more generally including those in overseas Instituts Pasteur, which clearly kept away from theoretical discussion concerning the origin of variations and adaptation in insects of medical interest.
Key words: medical entomology, Lamarckism, adaptation, biological races, Roubaud, Institut Pasteur.
Medical entomology took shape in France around
1900 in two main groups of institutions, one dominated by the Faculté de médecine of Paris and its
annex, the Institut de médecine coloniale, and a second dominated by the Institut Pasteur and its subsidiaries in the French colonies and protectorates,
soon to be associated with the French Army (Navy)
tropical health services (Opinel, 2008a). Medical
entomology emerged as a semi-autonomous field, a
compulsory associate of parasitology, alongside classical entomology which was mostly centred in the
Museum national d’histoire naturelle in Paris from
the beginning of the 19th century. The overlap
between the three groups of researchers was small,
and was in part ensured by the participation of most
of them in the Société française d’entomologie
(Cambefort, 2008) and the Société de pathologie
exotique after 1907 (Opinel, 2008a).
To a certain extent, the discovery that microscopic organisms were responsible for parasitic diseases
was not markedly different from the discovery that
microbial agents caused familiar infectious diseases.
But their transmission remained elusive for years.
The description of the contribution of insects to the
process of infection had been a scientific and medical breakthrough particularly in offering the possibility of interrupting, at some point, the triangular
relation between insect, parasite and man, as
expressed by Grassi’s law on malaria (Fantini,
1994). Field reality in medical entomology immediately proved much more complex than for microbial
diseases: physicians and scientists were faced by a
multi-faceted landscape of variations in the biological features of insects and parasites, including
changes in morphology of the latter during their
cycle, as well as multiple symptoms of diseases associated with identical agents (da Silva, 2005). In two
Correspondence: Gabriel Gachelin, UMR, 7596 CNRS - Université Denis Diderot, 54 rue de Picpus, 75015 Paris, France,
e-mail: ggachel@club-internet.fr
different geographical locations, local “adaptation”
of seemingly identical insects was demonstrated by
significant biological differences (Opinel, 2008b).
Thus, the description of the geographical distribution of variations and the understanding of their origin had become, as early as on 1900, key issues in
medical entomology and parasitology.
The Facultés des sciences where zoology, and
hence entomology, was taught to students and to
future researchers and teachers, played little part in
the cosmos of French medical entomology; the latter appeared more as the private property of physicians and veterinarians. On the other hand, the Facultés des sciences were better prepared to approach
problems raised by the identification of variations
within species and the description of complex biological cycles of organisms. It happened that questions of variability and adaptation in medical entomology were asked at the moment when general
biology and theories of evolution, themselves aimed
at understanding variability, were gaining chromosomes as the physical support in favour of Darwinism and selection processes. This position was, however, rejected by most French biologists, who argued
against Darwin and Weissmann (Tort, 1996). Natural sciences, zoology, botany in France, were
engaged in an altogether different, original approach
which resulted at the end of the 19th century in the
formulation of theories of transmission of phenotypic characters and adaptation of organisms based
on nutrition processes sensu lato. This approach
largely excluded chromosomal genetics. These theories can be grouped under the denomination of neoLamarckism.
The founders of French medical entomology were
trained in that context and were largely exposed to
that particular, not to say national, view of heredity
and adaptation. The present paper is an attempt to
delineate the influence which French theories of
genetics and evolution at the turn of the 20th century may have had on medical entomology. In par-
268
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G. Gachelin and A. Opinel - Influence of Lamarckism on Medical Entomology in France
ticular, it offers perspectives into the scientific
approach to variability developed by the prominent
French entomologist, Emile Roubaud (Opinel,
2008b).
A rift between French biology and
chromosomal genetics and Darwinian selection 1
Following the elucidation of the determinant role of
the nucleus by Weissmann in 1883, two successive
groups of discoveries gave heredity the support it
lacked: Mendelian genetics was rediscovered in
insects, plants and mammals, and the chromosomal
inheritance of characteristics was proven. This led to
a radicalization of attitudes towards theories of evolution in the period 1900-1930. Emphasis was either
placed on the selection of genetically determined
traits, a distinctive feature of Darwinism, or on a
progressive adaptation to environment, a distinctive
trait of neo-Lamarckism, a set of theories built up
after Darwin’s publications in an attempt to translate
into scientific concepts the Lamarckian philosophical notion of hérédité des caractères acquis. Darwinism implied discontinuous events such as mutations and the stability through generations of the
traits on which selection acted, whereas neo-Lamarckism postulated the existence of continuous and
rapid adaptations induced by and in response to,
changes in the environment. Thus a relative instability of the considered trait was assumed, which only
ultimately might become stably acquired. Were the
signs of adaptation of insects described by medical
entomologists (Opinel, 2008b) hereditary and resulting from Darwinian selection, or were they unstable
features induced by the environment and only
becoming gradually acquired, as postulated by
Lamarckian scientists? Should the latter hypothesis
be proven, it would in turn become possible to influence insect characteristics by suitably altering the
environment.
The introduction of evolutionary thinking in biology revealed a fault-line separating French biologists, who predominantly adopted neo-Lamarckism,
from biologists elsewhere who generally accepted
Darwinism. The singularities of French genetics and
approaches to evolution have extensively been discussed elsewhere (Burian and Gayon, 1999). Their
history can be divided into three periods: pre Weissmann’s establishment of the domination of the cell
nucleus on heredity around 1883; from 1883 to
1920, a period characterized by attempts to prove
experimentally Lamarckism; and post 1920-25,
which witnessed the progressive renewal of genetics
in France (Burian and Gayon, 1999) despite the fact
that Lamarckism long remained the commonly
accepted interpretative framework for evolution and
adaptation in France (Corsi, 1997).
1
This paragraph largely rests on the studies made by:
Thomas M. (2004); La Vergata A. (1996); J.-L. Fischer and
W. Schneider (ed.) (1990); R.M. Burian and J. Gayon (1999).
Lamarck’s proposals for a theory of transformation that linked different species in a time sequence
was associated with the hypothesis that transformation of species was based on the hérédité des caractères acquis (further “heritability of acquired characteristics”) induced by changes in environment and
by vital requirements, were enunciated at the beginning of the 19th century (Lamarck, 1801). Darwin’s
book On the origin of species by means of selection
or the preservation of favoured races in the struggle
for life, was translated into French soon after publication in 1859, and was widely read and discussed.
Darwin’s influence extended far beyond scientific
circles and engaged French intellectual life in general. Emile Zola offers one example of that trend
(Brown, 1996), and is particularly interesting in that
he amalgamated elements of Darwin’s and Lamarck’s proposals along with Prosper Lucas’ (18051885) natural heredity (Lucas, 1850). Such confusion was common in France, Darwin being considered as having merely reformulated Lamarck’s transformism. The depth of the difference existing
between gradual transformism and selection-driven
evolution had, in fact, not been well recognised
(Jacob, 1970) and the philosophical dimensions of
Lamarck’s proposals were considered scientific,
including by informed scientific circles as shown by
the talks delivered much later by Edm. Perrier and
Y. Delage, both professors at the Faculté des sciences of Paris, on the occasion of the dedication of
Lamarck’s statue at the Museum national d’histoire
naturelle in 1909 (Perrier, 1909; Delage, 1909).
The second period stemmed from the introduction
in 1883 of the notion of germ cells by August Weismann (1834-1914), which very plainly ruled out
and rendered scientifically unacceptable, the theory
of the heritability of acquired characteristics. That
was not easily accepted. The re-formulation of
Lamarck’s principles into a scientific theory opposable to Darwin, named neo-Lamarckism, was first
developed in the USA2, and was popularized in
France around 1885-1890 by the influential zoologist Alfred Giard (1846-1908) (Bouissy, 2004). The
rediscovery and the translation into French of
Mendel’s laws in 1900 was soon followed by the
enunciation of the chromosomal theory of heredity
first proposed by Walter S. Sutton (1876-1916) in
1902. This destroyed any remaining possibility of
syncretism between Darwinism and Lamarckism:
the transmission of phenotypic traits through the
transmission of chromosomes negated any direct
influence of environment or behaviour on the genetic material of germ cells, and thus excluded the possibility of a gradual and unstable acquisition of characters through generations by experience or necessi2
The expression “neo-Lamarkism” has been created in the
USA by the entomologist Alpheus S. Packard (1830-1905) in
1885. He was the author of a book entitled Lamarck, the
Founder of Evolution: His Life and Work, Longmans Green,
New York, 1901.
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ty. Despite the accumulation of scientific data on
chromosomal inheritance and on mutations as
source of variation, the belief in the heritability of
acquired characteristics, or at best a syncretic attitude, persisted among the majority of French biologists. Rejection of Mendelian genetics and the long
lasting existence of neo-Lamarckism were not due to
ignorance. French biologists were well informed
about Mendelian genetics. A vigorous debate was
going on between the proponents of Lamarckism
(such as Félix Le Dantec (1869-1917) and Giard at
the Faculté des Sciences de Paris), and opponents
like Lucien Cuénot (1866-1941) at the Faculté des
sciences de Nancy and seed producers such as the
Vilmorin family. Nevertheless, rejection of chromosomal inheritance was the common rule. Genetics
was not taught until the late 30s, and Lamarckism
remained the dominant philosophy taught in universities at least until 1950.
The surprising longevity of this situation in France
cannot be explained by scientific, but rather by
deeply rooted cultural and philosophical, factors.
The pre-existence of Lamarck before Darwin, and
the widespread acceptance of the heritability of
acquired characteristics, had made Lamarckism part
of the French cultural heritage. Lamarckian views
also encapsulated an understanding of heredity not
seen elsewhere but which prevailed in French biology. The French intellectual tradition in physiology in
fact appears to have played a major role in the widespread acceptance of Lamarckism by scientists. In
the middle of the 19th century, Lamarck’s philosophical transformism gained support from the
physiological views on heredity expressed by Claude
Bernard. Burian, and Gayon (1999) pointed out that
heredity in France was viewed, largely following
Claude Bernard’s speculations, as a particular part
of the general nutrition processes of organisms:
nutrition processes determined the mechanisms of
inscription and the transmission of characteristics:
all questions relevant to heredity converged to questions of nutrition at the cell level. Gayon, discussing
Bernard’s theory, concluded that, for Bernard, variations caused by environmental, physiological and
nutritional changes, might somehow be transmitted
to descendants. He notes the “extraordinary persistence of the thesis according to which heredity is a
facet, among others, of assimilation” (Gayon,
1991). That view, developed by theoreticians of neoLamarckism, particularly Le Dantec (1909), contributed to the speculative approach to genetics that
persisted in France throughout the first third of the
20th century, and to a certain extent until 19503.
3 The debate took a political aspect after WWII following
the Congress of the Academy of the Agricultural Sciences of
the USSR, hold in Moscow in 1948 which saw the triumph of
Lysenko. At that moment, some prominent French biologists
were either members of the Communist Party or were close to
it. They were asked by the Party to accept proletarian genetics and reject the classical, morganian approach oh heredity.
269
Some Lamarckian scientists developed their theory
out of their own field studies and tried to accommodate Darwinian components. Giard accepted the
existence of mutations (Thomas, 2004) 4 , but
thought they always had been silently effected by
exposure to changes in the environment and played
a minor role in evolution (Giard, 1904). He clearly
distinguished primary causes of evolution (some
being climatic in nature, such as temperature or
wetness, others being physiological in nature – food,
general biology, behaviour) from the secondary
causes which included heredity. Giard’s approach
could be considered as synthetic, since the role of
heredity was not ignored but only played down.
Other influential biologists raised more philosophical and radical objections to Mendelian genetics and
Darwinism: for Le Dantec, another theoretician of
biology, Weismann’s and Morgan’s approaches to
heredity were metaphysical and considered as reintroducing pre-formation theories and giving too
much importance to the nucleus compared to the
cytoplasm, though the latter was the very heart of
nutrition processes at the cell level and the basis of
heredity. In Evolution individuelle et hérédité published in 1898, Le Dantec developed a Bernardinspired law which he named loi de l’assimilation
fonctionnelle as the basis for evolution. In so doing,
Le Dantec abandoned experimental science to deepen his theoretical approaches to life. For him, heredity was decidedly a nutritional issue, and cytoplasm
played the dominant role 5. Mendelian genes should
be considered as microbes and variation as the
product of infection. Genetics and Darwinism
should thus be rejected: “on peut établir en partant
des seuls faits d’observation que j’ai signalés (multiplication et variation) les deux principes de
Lamarck, celui de l’adaptation et celui de l’hérédité
des caractères acquis” (Le Dantec, 1904). Etienne
This was the origin of a genuine drama inside the community
of French (not only) biologists. As an example of the continuation of Lamarckian thinking adapted to genetics, a noted
biologist Marcel Prenant, wrote in La Pensée (the philosophical journal of the French Communist Party) “la génétique
classique a glissé sur une pente dangereuse et réactionnaire,
que trop souvent le mendélisme moderne, morganien, a incorporé des idées ridicules sur l’indépendance des cellules germinales par rapport au milieu” before being himself excluded
from the Communist Party. M. Prenant “Un débat scientifique
en Union Soviétique”, La Pensée 1948, 21: 29-32.
4
Quoted from M. Thomas (2004): for Giard, she says,
mutations are “the brutal and sudden occurrence of a character that did not exist before, but which may have been prepared very slowly in the ancestors of the individuals it
appears”.
5
“Il faut concevoir que chaque espèce vivante a son protoplasme particulier… et la forme vivante est nécessairement
liée à cette composition protoplasmique: dans le milieu où elle
est adaptée à vivre, c’est-à-dire à assimiler, à accroître sa
masse, une substance vivante prend nécessairement la forme
spécifique correspondante. Et l’hérédité elle-même qui semble
si souvent mystérieuse, se réduit à la transmission directe de
cette substance spécifique des parents à l’œuf et par le l’œuf
au nouvel individu”. Le Dantec, quoted by Perez (1917).
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Rabaud (1868-1956), professor at the Faculté des
sciences, was also a radical but in a different way,
attacking less Darwinism than the chromosomal
theory of heredity. E. Rabaud considered that an
organism is nothing without its environment, does
not even exist since organism and environment are
two parts of the same whole; variation is the product of the interaction of the organism with its
changing environment. Rabaud, founder of French
ethology, is better known for his Lamarckian positions in medical anthropology and psychology
(Rabaud, 1908) 6. His studies on insect behaviour,
on line with popular Fabre’s descriptions, were the
basis for a Lamarckian approach to ethology, a dominant perspective until World War II. As a theoretician of the influence of the environment on behaviour, he contributed to the development of the
French school of experimental psychology, which
largely used insects as models (Thomas, 2003;
Dupont, 2005). After Giard’s death, he gave his
journal, the Bulletin scientifique de la France et de
la Belgique, a strong anti-Mendelian orientation.
Anti-Mendelian views were strongly challenged in
France after 1925. Lamarckism was gradually dissociated from anti-Mendelianism and lasted longer
(Corsi, 1997) in the context of a society rather unfamiliar with selection, in contrast to the Anglo-Saxon world (Burian and Gayon, 1999). Lamarckian
views of adaptation to environment sounded as an
obvious explanatory system by providing an intuitively understandable mechanism. In that respect,
the questioning of adaptation raised by field observations of medical entomology, which involved multiple interacting parameters resulting in complex
characters, appeared not less intelligible than
Lamarck’s own examples proposed as primum
movens of gradual change leading to biological
races, then to species. In an opposite way, adaptation to complex climatic features, or the elaboration
of behaviours, were roadblocks for French Darwinists since the selection of phenotypical traits by complex interacting features could not be explained by
mutational theory as easily as simple geneticallydetermined characters were supposed to be: mutationism was culturally difficult to admit as a driving
force for macroevolution.
Thus, the “French style” in genetics and evolution
of the period 1900-1930, was dominated by the paradigm that progressive trans-generational phenotypic adaptation of organisms to environmental and
nutritional changes would finally lead to the inheritance of characteristics gradually acquired during
adaptation processes. That conclusion did not mean
that French biologists were all active proselytes of
neo-Lamarckism. It merely indicates that they were
taught by, and worked with, a few dominant zealot
6
His attacks against Lombroso and his notion of “criminal
né” remains frequently quoted. See Rabaud E. (1908), Le
génie et les théories de M. Lombroso, Mercure de France,
Paris.
neo-Lamarckian scientists at the university and in
marine laboratories, and thus could have been
inclined to consider Lamarckism as the only valid
theory of evolution. Nor did they receive any training in genetics. The diversity of opinions emerging
after 1925 forbids consideration of the existence of
a monolithic Lamarckian school of evolution in
France, but Lamarckism was simply defined as
belief in the heritability of acquired characteristics,
kept functioning as a cultural consensus in France
(Corsi, 1997), well evidenced by its teaching in secondary schools until the end of the 1950s, and
covertly into the present 7.
Could Lamarckian scientists have influenced
medical entomology?
The above conclusion does not imply that Lamarckian biologists exerted a significant influence on medical entomologists. The community of medical entomologists was comparatively small and developed
out of mainstream entomological research, particularly where physicians were concerned. By contrast,
parasitology, with which medical entomology was
closely associated, was an active domain of research,
and one in which fundamental biology had been
interested in since the middle of the 19th century:
among the first demonstrations that a parasitic
worm was transmitted by insects was that of Henri
de Lacaze-Duthiers (1821-1901), a zoologist pioneer of the creation of marine laboratories, in his
1853 study of plant galls (Lacaze-Duthiers, 1853)8.
The ability of a physician-parasitologist like Emile
Brumpt (1877-1951) to elucidate complex parasite
cycles was largely due, he claimed, to his zoological
training in marine laboratories. He was rather proud
to write that he worked as a zoologist (Opinel and
Gachelin, 2004), an example of the absence of barriers between Faculté des sciences and Facultés de
médecine in the present domain of interest.
7
Lamarckian wording persists up to now. In a recent issue
of Biologie Géologie, the journal of high school teachers of
natural sciences, we quoted, in an otherwise neutral article on
liquid food, the following sentence “Pour pouvoir s’alimenter
et perpétuer l’espèce, les Endoparasites calquent leur cycle
biologique sur l’activité nychtémérale de l’hôte”, a sentence
which implies that the parasite has deliberately chosen to use
the distinctive features of the host for an efficient reproductive cycle. In the same article, differences in the structure of
the buccal apparatus of hymenopters are presented as an evolutionary series. Clos J. (2007), L’alimentation liquide chez les
animaux, Biologie-Géologie, 4: 709-758.
8
Lacaze-Duthiers H. de (1853), Histoire des galles. Ann.
Sc. Nat. Bot. 3ème series, tome 19, page 273. Quoted by Darwin in The Variation of Animals and Plants under Domestication. Lacaze-Duthiers had a considerable influence on
French zoology through researches on marine invertebrates
carried out in two marine laboratories he created, Roscoff and
Banyuls: microscopic observation of micro-fauna and their
development, description of local “ecosystems”, description of
parasitic and symbiotic lives, were among the most frequent
studies carried out in these laboratories and nearly all French
parasitologists got part of their training there.
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Let us first examine the place given to Lamarckism in the principal (Parisian) research and teaching
French institutions dealing with entomology and
medical entomology. At the turn of the 20th century, dipterology (to which medical entomology could
at that time somehow be reduced), was a minor
activity at the Museum national d’histoire naturelle.
Emile Bouvier (1856-1944), president of the Société
française d’entomologie in 1898, re-introduced
dipterology after 1904 by first hiring the dipterologist J. Villeneuve de Janti. From that moment on,
lectures on flies of medical interest given at the
museum were considered as companion lectures to
the main parasitology course given at the Institut
Pasteur by Laveran and Mesnil. Bouvier’s opinions
concerning the evolution of insects were clearly on
the Lamarckian side, particularly concerning the
progressive acquisition of food behaviours (Bouvier,
1918). Emile Roubaud (1882-1962) a future prominent medical entomologist was introduced to
dipterology by Villeneuve in 1904 in Bouvier’s laboratory. Bouvier participated in Roubaud’s nomination as the zoologist to the mission on sleeping sickness in the Congo, which determined nearly all
Roubaud’s subsequent career (Opinel, 2008b). Bouvier was succeeded in 1931 by René Jeannel (18791965), a specialist in subterranean coleopterans,
who worked on the morphology and adaptation of
their sex organs. President of the Société française
d’entomologie in 1932, Jeannel was convinced that
the adaptive characters of these organs, as well as
the reproductive habits of coleopterans, had been
slowly and gradually acquired for the realization of
their function, to progressively become hereditary
(Urbain, 1946). Lamarckism was thus still the
explanatory paradigm of evolution for classical entomologists before World War II. It is interesting to
note that Jeannel extended and largely popularized
the earlier opinions of Jacques-Henri Fabre, according to whom Darwinism could not explain instincts
and behaviours. This approach also fited well with
Roubaud’s publications on insect ethology, and, in a
general way with – once again – a distinctive French
approach to ethology which incorporated Lamarckism in its explanatory systems (Thomas, 2004).
If we now turn to the university, Lamarckism
there was a dogma, particularly at the Faculté des
sciences of Paris. Lamarckian scientists occupied
two strongholds in Paris, located at the Faculté des
sciences, and at the Ecole normale supérieure,
respectively. They were not physicians but zoologists
(dominantly malacologists, protistologists and entomologists, usually experts in all fields of zoology) or
botanists, and were renowned for their contribution
to the taxonomy and physiology of plants and animals, including parasitic and symbiotic relations
often considered as a theoretical basis for biology
(Caullery, 1922). In general, their involvement with
evolutionary theories appears less rooted in their
scientific work than in their philosophical opinions
about life, biology and philosophy. The two main
271
theoreticians of biology and evolution, who dominated the years 1890-1914, were also university professors in Paris. It is beyond doubt that Alfred Giard (1846-1908) occupied a dominant, not to say
overwhelming position in research and university
teaching in all fields of invertebrate zoology from
protistology to entomology. A convinced Lamarckian scientist, he effected the creation of the Laboratoire d’évolution des êtres organises at the Sorbonne
in 1888, where he taught neo-Lamarckism until his
death in 1908. He also was director of the Lamarckian journal Bulletin scientifique de la France et de
la Belgique from 1878 9, and founded the Laboratoire de biologie marine in Wimereux, which aimed
to train zoology students in marine invertebrate
biology. His influence was also extended by his institutional positions: president of the Société entomologique de France in 1896, president of the
Société de biologie in 1904 and president of the
Société française pour l’avancement des sciences in
1905. In 1908 he became a member of the board of
governors of the Société de pathologie exotique. He
was elected at the Académie des sciences in 1900.
As regards medical entomology stricto sensu, he was
an active member of the committee which organised
Roubaud’s mission to study sleeping sickness and
the biology of Glossina sp. in Western Africa. and
he was in charge of writing the zoological instructions for the mission (Bouvier, Giard and Laveran,
1906). Giard’s laboratoire d’évolution des êtres
organises, in addition to being a breeding ground
for young biologists, was dominated by Lamarckism
until 1955, a remarkable longevity. Among the many
members of Giard’s laboratory, Roubaud has already
been mentioned. As the founder of French insect
ethology Giard was twice president of the Societé
française d’entomologie, in 1916 and 1923. Maurice
Caullery (1868-1958) developed a more ambiguous
and complex intellectual position concerning evolution theories. A renowned zoologist, specialist in
protists and marine invertebrates, particularly parasites, he first introduced Darwinism to France before
adhering to the heredity of acquired traits when joining Giard’s group around 1905. Later, after World
War I, he developed a personal, rather theoretical
approach to evolution of organisms. Caullery succeeded to all the positions Giard had occupied.
Through his books and lectures on transformism, he
exerted a strong influence on French biology until
1950 at least. As for Giard, he never rejected either
Darwinism nor mutations but considered them as
minor causes of evolution. Caullery was succeeded
by P.-P. Grassé (1895-1985), president of the
Société française d’entomologie in 1941 and himself
9
The Bulletin scientifique de la France et de la Belgique
was the journal in which Lamarckian views on evolution were
published. Its editorial board contained most of the persons
mentioned in the present section, who could thus not have
been opposed to Lamarckian views on evolution, due to the
personalities of Giard, Le Dantec and Rabaud.
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rather on the Lamarckian side concerning animal
and insect behaviours. The reasons for the persistence within the Faculté des sciences of Paris of a
large research and teaching institute dominated by
Lamarckism until at least 1955, remain to be studied10. The second important theoretician of Lamarckism was Le Dantec (1869-1917), mentioned
above, who had strong links with Giard. He played
no role in the teaching of entomology, but taught
embryology and general biology to students at the
Faculté des sciences and at the Ecole normale
supérieure. He was primarily a “scientific philosopher” of life, expressing his opinions in several
books largely inspired by positivism and Lamarckism (Perez, 1917). Le Dantec exerted a significant
influence on political, rationalist and positivist circles as a kind of “theologian of scientism” (Balibar,
2000) and on students through his teaching duties
in embryology and general biology until his death in
1917. He does not, however, appear to have created a distinctive school of thought.
The situation of the Institut Pasteur and the overseas Pasteur institutes, where medical entomology
developed in the laboratory and was used in field
research and prophylaxis, differed profoundly from
that of the Faculté des sciences. Indeed, research
carried out at the institute was intended to result in
improvements in public health. In that respect, the
recruitment of parasitologists and entomologists to
the Institut Pasteur (Paris) could appear anomalous
since it involved zoologists who were not physicians. Recruitment for the institutes abroad was dictated by field efficiency and took place principally
among military physicians (Opinel, 2008a). However, Felix Mesnil (1868-1938), a biologist devoid of
medical training, introduced the study of tropical
diseases and medical entomology as early as 1898.
His opinions concerning evolutionary theory are not
known with certainty. The titles of the papers he
selected for analysis in the Bulletin de l’Institut Pasteur from 1903 (signed FM) reveal his interests as
nearly exclusively in parasitology and tropical disease. He also worked on marine invertebrates with
Caullery, his brother-in-law. He became a member of
the editorial board of Giard’s journal in 1909, which
implies that he adhered to the trends oriented
against genetics and evolution. In contrast to the
laboratoire d’évolution des êtres organises, there
was no collective position at the Institut Pasteur
concerning evolutionary theory, though Burian and
Gayon (1999) suggest that Larmarckism was present in several laboratories. The intellectual atmosphere of Mesnil’s laboratory can be approached
through the researches that were carried out there.
After 1920, E. Chatton and A. Lwoff were studying
the inheritance of external features of ciliates and
The imposant building of the laboratory, located 105
boulevard Raspail in Paris, is not anymore a laboratory and
houses part of the Ecole des hautes études en sciences
sociales.
10
the influence of nutrition, with the idea of a non
chromosomal inheritance and a Lamarckian evolution of the morphological features of ciliates (Burian and Gayon, 1991). The famous expression
“adaptation enzymatique”, the study of which was
launched in Mesnil’s laboratory, is in itself rather
ambiguous in view of the meaning “adaptation” had
in the context of French biology before World War
II. Mesnil’s laboratory was thus most probably not
hostile to Lamarckism. As regards medical entomology per se, Mesnil also pushed Roubaud, a member
of his staff, to participate in the mission on sleeping
sickness in Western Africa. Roubaud’s scientific
work on the role of environment in adaptation
processes and biological races is discussed in the
accompanying paper (Opinel 2008 b). The strong
influence of Lamarckian thinking on Roubaud’s theories is presented later in the present paper.
By contrast, the influence of Lamarckism appears
to have been weak or absent among most of the
physicians dealing with arthropod-borne diseases,
medical entomology and parasitology, whether at
the Faculty of medicine or in colonial institutions.
Medical zoology, soon followed by parasitology, had
been extensively taught since 1883 at the Faculté de
médecine of Paris by Raphaël Blanchard. The teaching of medical entomology was introduced at the
faculty of medicine as soon as the notion of the vector was defined (Opinel, 2008a; Osborne, 2008).
Blanchard was succeeded by Emile Brumpt and coworkers such as Neveu-Lemaire and Langeron. A
genuine hot spot of research and teaching on parasitology and entomology (as was the military medical school of the Pharo in Marseilles on the applied
side), the Faculté de médecine de Paris and its
annex, the Institut de médecine coloniale, were
from the beginning positioned well away from the
theoretical discussions on races and adaptation prevailing at the Laboratoire d’évolution des êtres
organisés at the Sorbonne and around Roubaud at
the Institut Pasteur. The survey of the Traité de
pathologie exotique of Rist and Jeanselme (1909)
shows it to be entirely devoted to diagnosis, treatment and prevention of these diseases. A survey of
successive editions of Brumpt’s treatise shows that
the author used the word “adaptation” but did not
give it more than its usual meaning 11. A purely practical approach to medical entomology was also
taught at the Institut de médecine coloniale annex
of the Faculty of medicine and at the Navy application school of Le Pharo. An exception may perhaps
be found in the treatise on tropical diseases by
Alphonse Le Dantec (1909), which was introduced
by chapters inspired by the French medical geography of the 19th century ,and involved a theory of
11 For example, in Brumpt, edn 1922 p. 15: concerning
morphological and biological adaptation of parasites: tous ces
phenomènes sont le plus souvent des adaptations favorables
à la conservation des espèces et leur caractère actuel a dù se
sévelopper progressivement.
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adaptation to climates (Gachelin, 2005). The factual, neutral, approach adopted by physicians, and the
absence of theoretical developments, does not imply
a lack of interest in the complexity of the systems
they dealt with. On the contrary, they wrote extensive descriptions of the habitats and environments
of insects with a view to designing effective preventive strategies. The parameters they used to screen
environmental effects included insect behaviour
(blood meal preferences, site of reproduction, flight
length) but placed the emphasis on those which
could interfere with development (eggs laying and
conditions of larvae survival), on precise taxonomy
and on the association with microbes. On the
whole, physicians described the characteristics of
infestation and insects more than they indulged in
biological interpretation of the adaptations to environment which they noted. Such a conclusion
extends to important missions carried out before
World War I, often under the supervision of the
Institut Pasteur. Marchoux and Simond’s mission to
Rio de Janeiro to study the transmission of yellow
fever resulted in precise description of the habitat
and biting behaviour of an insect (Lowy, 2001) but
with no evolutionary interpretation. In their works
on malaria and other diseases, and on insect vectors,
Et. and Em. Sergent in Algeria (Sergent, 1962)
studied the behaviour and biology of anopheles
extensively, but in order to identify them and their
habitat, for the better calculation of ways to combat
the insects. Similarly, Brumpt’s 1903 mission to the
Congo concerning sleeping sickness resulted in suggestions for the best manner to fell woods on river
banks to reduce contact between human populations and glossina (Brumpt, 1903). The 1924-1931
Brumpt mission to Corsica reported very accurate
descriptions of the ecosystem of the local anopheles,
but primarily as a way to define sites for preventive
action (Opinel and Gachelin, 2004).
It can thus tentatively be concluded that Lamarckism was deeply influential in the Faculté des sciences, was part of the scientific atmosphere of the
Institut Pasteur (Paris) and of the Museum national d’histoire naturelle but was not adopted by physicians. It was a genuine paradigm for university zoologists and entomologists, but was at best interpreted by physicians as an “easy explanation” for those
working on field medical entomology.
An experimental approach to
the Lamarckian view of adaptation
The nature of food, climatic conditions, physical
and environmental traumas, behaviour and even a
kind of intentionality of the organism in its search
for the proper food, the proper climate or the best
reproductive conditions so as to properly adapt to
them, were thus assumed to be the driving forces of
variability and adaptation according to Lamarckian
views on evolution. These assumptions were
strengthened by the notion that an organism cannot
273
be defined independently of its environment. Food,
behaviour and climatic changes were all embedded
in the environment, as also were the features that
insects of medical interest had to adapt to (Opinel,
2008b).
Lamarckism has also been tested in experimental
approaches either aimed at proving the hereditary
influence of environment on the evolution of
species, or as a tool to dictate the orientation of
species towards different biological characteristics.
Darwinian selection theory was beyond any experimental approach. On the contrary, the progressive
accumulation of discrete adaptive changes was
thought to occur rapidly and believed to operate on
a shorter time scale, and thus was amenable to an
experimental approach. Between 1885 and 1920,
several French biologists attempted experimentally
to verify the heritability of acquired characteristics
by introducing, during a number of generations,
repeated changes in the conditions of life and in the
food of animals and plants.. some typical experiments which bring us close to these experimental
attempts at inducing changes in species.
Changes could be traumatic. Louis Blaringhem
(1978-1956) carried out repeated mutilation of the
reproductive organs of maize plants to induce a significant enough alteration in their physiology that
the mutilation could become imprinted in the whole
organism, including the reproductive cells (Blaringhem, 1907).
Changes could directly affect the nutrition processes of the whole organism and result in adapted characteristics: Blaringhem altered the nutritional support of maize grown in open fields. He concluded
that new characteristics could result from nutritional change but they were not stable. The finding that
entirely new strains appeared that were immediately genetically stable, led Blaringhem to move closer
to Darwinism (Thomas, 2004).
Changes could be climatic and lead to adapted
characteristics: Gaston Bonnier (1853-1922) a
botanist still known for his popular flora, carried
out long term experiments to adapt plants in his laboratory at Fontainebleau or in alpine conditions, to
different climates and to create newly adapted races
of plants, by growing and seeding them over a 30
year period under climatic conditions different from
those prevailing in their original habitat (e.g. from
low to high altitudes, which includes a number of
climatic parameters).
Changes could be directed at creating mutations.
Of primary interest in the context of the present
study, since the experiments were carried out on
flies, Etienne Rabaud, Alfred Giard and Maurice
Caullery, faced with the growing importance of the
mutation theory, asked a PhD student, Emile
Guyenot (1885-1963), to determine the mutation
rates of Drosophila melanogaster flies fed over several generations on chemically different nutrients.
Guyenot’s supervisors expected that differences in
food would result in different mutation rates. The
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experiment was important since it well formulated.
After studying 400,000 flies and their offspring, and
having noted a large number of mutations, Guyenot
concluded that food composition had no effect on
mutation rate (Burian and Gayon, 1999). The publication of his thesis in 1917 created furore in the
Laboratoire d’evolution des êtres organisés, and
affected Guyenot’s career: he was unable to find a
position in France and became professor of biology
at the university of Geneva12, where he specialized
in teratology and in 1924 wrote the first textbook
on genetics to be published in French.
Experiments were thus actively carried out in
France at least until World War I (the experimental approach appears not to have been resumed
after the war) to prove the heritability of adaptive characteristics acquired through changes in
food and climatic environment, and through trauma. It is remarkable that neither negative results nor
research progressing elsewhere in genetics altered
the general French consensus that heritability of
acquired characteristics explained variations and
evolution.
Few professional, university-trained entomologists
had in fact worked in medical entomology or studied the adaptation of several different vectors and
microbes in the period 1900-1940. In France, Emile
Roubaud has been one of the few to have approached problems of medical entomology as a genuine entomologist, and not as a physician obliged to
deal with insects vectors of disease. Roubaud is of
further interest in that he studied the biology of
races of several insect species along similar lines and
tried to uncover general laws of biological adaptation in insects (Roubaud, 1935). The manner in
which he conducted observations and experiments
on Glossina and Anopheles, and the conclusions he
drew from them, are presented in the accompanying
paper (Opinel, 2008b). We discuss here them in relation to the heritability of acquired characteristics.
As a student and a young researcher, it should be
emphasised, Roubaud was located in the context of
triumphant Lamarckism. He was trained at the Faculté des sciences of Paris, became Licencié de sciences naturelles in 1901, and Agrégé de sciences
naturelles in 1904. He was taught by Giard at university and by Le Dantec and the Lamarckian
botanists during his preparation for the Agrégation
given at the Ecole normale supérieure. He was further trained in dipterology at the Museum national
d’histoire naturelle. In the mean time he attended
lectures given by Giard at the laboratoire
d’évolution des êtres organisés. The conclusion that
he had been infused with Lamarckian ideas on the
origin of evolutionary variations appears difficult to
avoid. Did this intellectual position have any effect
on his collection and interpretation of data? There
are no direct references to Darwinism or Lamarck12 That anecdote well illustrates the power of Lamarckian
scientists in French universities.
ism in Roubaud’s documents and Titres et travaux
examined. Roubaud developed his own methodical
system of interpretation concerning the influence of
the environment on the adaptation of biological
races of insects, and designed experiments which, in
view of the above, are strongly reminiscent of the
research, experimental work and theory conducted
in the laboratories of Giard, Rabaud and Caullery.
Roubaud’s main results were drawn from the two
West African missions to study the biology of
Glossina and trypanosomes, and could be summarized as suggesting a strong correlation of the biological features of the organisms with climatic parameters, and the introduction of a theory of prophylaxis against sleeping sickness by zootropic deviation. His research introduced the notion of biological races of Glossina and led him to experiment in
the modification of the infecting power of Glossina
by exposing them to an atmosphere similar to that
found in areas where the infectious power was low
(Opinel, 2008b). In other words, the experiment
was aimed at reproducing in the laboratory the
process of adaption to climatic conditions as it
occurred in nature, resulting in the creation of distinct races. From a methodological view point, the
link between environment and geographical distribution resulted from correlation, the adaptive interpretation of which led to a typical Lamarckian
experimental program.
The idea of “educating” insects to make them
innocuous remained a constant theme in Roubaud’s
work. Whatever species of insect considered, the
idea stemmed from the conclusion that the existence
of different races of insects was associated with climatic differences: thus according to Roubaud, the
repeated exposure of an insect race to a different
environment will lead to a newly adapted race. Climatic environment, including natural alternation of
phases of humidity and dryness, also plays a determining role in the biology of insects and in the stability of species which, he argued, need to be regenerated (i.e. restitution of the archetype) from time to
time (Roubaud, 1924). In search of general laws of
nature, Roubaud introduced several theoretical
notions on insect biology that were linked to the
cyclical alternation of temperature and dryness. He
distinguished biological types for all insects as being
defined by the climatic parameters they are adapted
to. The original features of adapted insects could,
according to his theory, be restored (reactived or
regenerated, reactivation or regeneration appears as
a kind of return to the original phenotype) by cold
(athermobiosis) but not by dryness, or by dryness
(anhydrobiosis) but not by exposure to cold. Thus,
the distribution of biological races within species
usually is in accordance with annual temperature
and humidity cycles. An important role was given to
the alternation of cold and dry phases as the originators of new races, and as requirements for the
regeneration of the considered species (Roubaud et
Colas-Belcour, 1933), the characters of which may
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have changed with time, thus for example converting an homodyname species into the original heterodyname one (Opinel, 2008b). Roubaud writes
about a locust, Schistocerca peregrina « je crois pouvoir déduire que les générations rapides obtenues
sans l’intervention de l’arrêt d’anhydrobiose perdent
le caractère gregaria pour acquérir le type dissocians
ou flaviventris” (Roubaud, 1933). A specially
equipped laboratory, the insectarium, was built at
the Institut Pasteur in 1930 to produce large quantities of insects (anopheles and phlebotomes) at various stages of their development. It was also the site
of experiments aimed at modifying their biological
properties by alterations in temperature and humidity. Roubaud tried to induce adaptation in locusts
(to reduce their ability to migrate) and in phlebotomes (to synchronize their development) by controlling temperature and humidity during their larval development, a procedure he named “education”
(Roubaud and Colas-Belcour, 1927). As regarded
populations of wild insects, Roubaud argued that
insects deliberately chose the climates which best
suited their needs for reactivation. Moreover, he
suggested that this conclusion could be extended to
all migrating animals: the choice of place to undergo reactivation might be the key to understanding
the distribution of living beings, and the suitability
of the chosen place being determined by hygrometry
and temperature cycles.
Roubaud’s earlier studies on the zoophilic deviation of Glossina led him to develop a particular
interest in the origin of nutritional and reproductive
behaviours, zoophilic preferences being one example in the study. He listed instincts and behaviours
as acquired characteristics. As described elsewhere
(Opinel, 2008b), he regarded the relative “appetite”
for the blood of cattle or humans as responsible for
the emergence of the biological races of Anopheles.
How was that differential appetite established and
later stabilized? First, Anopheles adapt to the microclimate offered by human and cattle shelters, which
is different from external conditions, thus allowing
the persistence of parasite and vector in areas where
they are not found outside. Under these conditions,
Anopheles came into close contact with cattle, and
the attenuation of thermal variations by human
housing allowed Anopheles to develop into
zoophilic races. The preferences of insects in the
animals they bite is established by recurrent exposure: “par suite de l’hérédité d’accoutumance au
regime, les générations successives d’insectes issus
d’un meme hôte tendent à constituer des races physiologiques”, a clearly Lamarckian statement. These
new races are associated with the “technical” adaptation of their biting appendages to the skin, characteristics of either man or cattle (adaptation
hémophage) Opinel (2008b). The whole process
was placed under the double control of human habitat and hydrological conditions (Roubaud, 1928),
“c’est ainsi grâce au facteur de régulation hydrologique que se constituent peu à peu les races zoo-
275
philes françaises”. This in fine substitutes the relation habitat/man/animal for man/marshes in the
propagation or disappearance of malaria by Anopheles (Roubaud, 1928).
The chemical nature of food supply also appeared
to be a determinant of the emergence of new biological races through the acquisition of distinctive behaviours, for example during the feeding of larvae. The
feeding conditions associated with a particular location, for example town houses vs rural farm, were for
Roubaud the origin of the biological races of Culex
identified around Toulouse (Roubaud, 1930). For
example, the adaptation to conditions of human life in
towns was accompanied by the development of
cesspools, which favoured the adaptation of larvae to
a food rich in ammonia with important consequences,
particularly on the requirement for ovary development
of blood meals (autogene13 vs non autogene races)
(Roubaud, 1932). The heated debate between
Roubaud and P. de Boissezon is a good example of the
discussion of Roubaud’s proposals. Boissezon (1934)
contested Roubaud’s idea of a distinction between rural biological races of Culex (hétérodyname and anautogène) and town-dwellers (homodyname and autogène). In Boissezon’s view, Roubaud had merely
observed the consequence of a differential breeding of
larvas thanks to the plasticity of the species: a food
rich in iron induces autogenesis, a condition reversible
within the extent of plasticity of the species. Plasticity
refers to reversible adaptability to environment,
whereas adaptation, according to Roubaud, refered to
the acquisition of biological features. The use of the
notion of plasticity threatened the notion of adaptation. Roubaud’s reply was brutal: Boissezon was ignorant of the methods by which biological races might
be identified properly. He should have known that
“l’autogène se différencie en effet foncièrement par sa
sténogamie franche appréciable en cubage inférieur à
1 litre, de la race hétérodyname anautogène qui est
eurygame” (Roubaud, 1934) in which a well defined
characteristic, here the need for a blood meal before
egg laying, is substituted by reproductive behaviour.
For Roubaud, plasticity14 did not exist in Culex and
races of Culex did (Roubaud, 1934). His attempts to
feed Culex larvae variously so as to induce different
egg laying behaviour in adults failed, “la nourriture
larvaire n’influe pas sur le développement de
l’autogénèse chez les races de Culex pipiens spécifiquement autogènes” (Roubaud, 1934): a conclusion
which should have been taken as meaning that autogenesis has a taxonomic character. This is suggested in
a previous Roubaud paper, though the word “genetically” used in the title does not refer to genetics but
13 “Autogène” defines the ability to lay eggs without a blood
meal.
14 It is interesting to note that the debate between plasticity and adaptation correlates well with the beginning of
research in physiological genetics which implies, in contrast
with Lamarckism, a functional relation between genes and
phenotypic characters.
276
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to transmission to progeny, whatever mechanism is
involved (Roubaud, 1930). By contrast, he considered
that the biting behaviour of Anopheles races could
still be influenced by the nature of the food provided
to the larvae. Roubaud reported that “les imagos
femelles issues de larves soumises au régime carné pur
manifestent plus tardivement et de façon moins active
leurs besoins de sang que les moustiques provenant de
larves astreintes à un régime végétal pur. Ceux-ci
manifestent une agressivité notablement plus précoce
et plus active” (Roubaud and Treillard, 1934). The
difference between Culex and Anopheles was
explained by assuming that Culex isolates had been
geographically isolated (Paris, Toulon, Alger), were
adapted to climatic (temperature and humidity) and
human parameters, and corresponded now to fixed
entities resistant to influence, in contrast to anopheles
(Roubaud, 1933).
A key issue was the understanding of the meaning
given to the notion of biological race. As did most
field medical entomologists, Roubaud defined geographically isolated and biologically defined sub populations of insects as “races” i.e. groups of individuals
that could not be distinguished from other members
of their species on a mere taxonomic basis, but were
defined as sharing a distinctive biological (for example temperature of development of the larvae) or
behavioural (such as feeding habits, zoophilic or
anthropic races of Anopheles, reproductive behaviours, etc.) properties. After World War I, Roubaud
extended the notion of biological race to other insect
vector species. He was among the first scientists clearly to enunciate the existence of distinct races of
Anopheles maculipennis – zoophilic vs undifferentiated races – based on the different feeding habits of isolates of otherwise morphologically undistinguishable
creatures. This notion was more or less accepted by
other medical entomologists. The use of the notion of
race remained scientifically valid until genetic analysis
allowed the segregation of different hereditary discrete
features around 1925 15, and finally the identification
of genetically defined species on that basis by the end
of the 1930s. In short, the notion of biological races
was gradually replaced by that of genetic variants
(varieties or strains) within a species, or by species,
depending on each particular case, in a Darwinian and
genetic scientific perspective. The point where
Roubaud diverged from other medical entomologists
was in stating that environmental conditions generate
the biological races. Roubaud accumulated data and
experiments to prove his point, whereas most contemporary medical entomologists actively sought
15
Genetic polymorphisms within races of mice were discovered around WWI when searching for the proper conditions to transfer tumors from mice to mice. A detailed account
of the transformation of “races” of mice into genetically
defined “strains” of mice in the frame of a genetic analysis of
graft rejection, is discussed in Gachelin G (2006). La construction de la souris idéale, in Les Organismes modèles dans
la recherche biomédicale. Sous la direction de G. Gachelin,
Presses universitaires de France.
hereditary taxonomic markers, if any, to define races.
This resulted in the solution given by Hackett and
Missiroli in 1935 to the paradox of anophelism without malaria, based on the identification of several varieties within the A. maculipennis complex (Hackett
and Missiroli, 1935). That conclusion was questioned
by Roubaud until at least 1939, as is evident in his
correspondence with Hackett and members of the
Rockefeller foundation in that year 16. Thus, it appears
that Roubaud kept to his definition of races of insects
at least until World War II, insect races being
described by the assumption of their evolutionary origin through the constraints of environment. Biological
insect races could be bred, stabilized by using conditions which mimicked the natural environment, but
reversion or conversion into other biological races was
possible by imposing permanent environmental
changes. Thus, for Roubaud, biological races became
hereditary only where the environment was kept constant for long enough. For example, long term exposure to cattle in a stable environment promoted the
differentiation of zoophilic races: “La stabilisation des
conditions de vie du moustique est l’un des plus puissants facteurs de sécurité de l’action déviatrice animale” (Roubaud, 1928), provided the nature of the
cattle involved remained constant. An insect race
could not be dissociated from its environment,
whether animal or climatic, a statement strongly reminiscent of the environment-organism duo proposed
by Rabaud and which excluded an independent study
of the organism. The emphasis placed on behaviour
rather than on taxonomical markers was strongly reminiscent of contemporary French ethology, which was
largely influenced by Lamarckian ideas (Dupont,
2004; Thomas, 2004).
Discussion
The research carried out on variation among insects
by Roubaud in the period extending from 1905 to
WWII, was in many respects influenced by Lamarckism, considered as a theory of biological variation.
Roubaud’s experiments in globally modifying insect
species were strongly reminiscent of those carried out
in Giard’s laboratory. The postulate of a direct link
between morphological features such as the maxillary
index (Opinel, 2008b) and adaptation to feeding on
16
Rockefeller Archives center. RF 6.1, series 1.1, Box 24,
Folder 276. Roubaud 1932-1939. In 1939, Roubaud asks for a
grant to visit Albania’s Rockefeller foundation laboratory to
solve the conflict between Hackett and Roubaud about the existence of races of Anopheles maculipennis (Hackett to Warren,
Rome Feb. 4, 1939 “for some time, Professor Roubaud of the
Pasteur Institute, Paris, has been publishing results that conflict
with results that L.W. Hackett and Bates has gotten with certain species of anophelines”. Roubaud contests them on the
basis that it could exist two different races of A. maculipennis
“one zoophilic and the other including man in its range of
hosts, memorandum No. 15 Hackett to Warren, March 31,
1939). No grant was given (Hackett to Roubaud, March 7,
1939) on the motive “that the conditions in Albania are unsatisfactory for his visit” (Warren to Sawyer, July 13, 1939).
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certain animals, elaboration of food and reproductive
behaviours by adaptation to chosen targets, possibility of “regeneration” of species by climatic relocation,
the importance granted to behaviour rather than to
taxonomic characters, were all placed in the context
of a gradual heritability of characteristics. Roubaud
did not refer to the nature of the genetic mechanisms
that might account for his observations. In all this,
Roubaud appears to belong to the Lamarckian heritage. His teaching position at the Institut Pasteur may
have contributed to embedding French medical entomology in Lamarckian interpretations in the years up
to World War II.
Yet, the period between 1930 and 1939 witnessed
the gradual undermining of the Lamarckian interpretation of the adaptation to environmental parameters
developed by Roubaud and based on the inductive
strength of environment parameters and the progressive acquisition of characteristics. Several factors
explain this loss of explanatory power among entomologists. As noted, complex characteristics were not
easily amenable to genetic analysis. Thus a Lamarckian-type interpretation of adaptation was more or less
shared by most entomologists but consensus changed
with the genetic determination of individual components, the sum of which contributed to complex characters. This occurred, say after 1925, in the context of
a general movement dominant in the Anglo-Saxon
world, characterized by the development of population genetics and the genetic analysis of polymorphisms, largely within the English school of genetics,
but which was initially marginal in France, emerging
there only after 1932 (Givernaud, 2000). Thus, individual characteristics contributing to complex ones
could have been selected. Moreoever, the importance
granted to ecosystems, a move initiated at the end of
the 19th century, made possible an ecological description of the multiple interactions leading to equilibria
between animal and vegetable populations in a
defined geographical area (Acot, 1988; Drouin,
1993). The combination of genetic and ecological
notions allowed Hackett and Missiroli to answer the
long pending question of “anophelism without malaria” in 1935 (Fantini, 1994). The progressive changes
leading to biological races postulated by the Lamarckian approach had to be replaced by the selection of
a variety of polymorphisms and by the equilibrium
between selected, polymorphic, populations. Moreoever, insect physiology, particularly concerning the
endocrine control of reproduction and development,
was also expanding rapidly and offered new mechanistic interpretations of instincts and habits concerning blood meals and food in general17. The contestation of Roubaud’s theories by Boissezon in 1934 was
an early example of a change in attitude among
French biologists towards Lamarckian and nonMendelian theories of races. Boissezon, an insect
17
It is worth noting that VB Wigglesworth, the discoverer
of the endocrine control of ovary functions in insects, was in
1930 a researcher in the department of medical entomology
at the London School of Hygiene and Tropical Medicine.
277
physiologist, had shown in respect of Culex in 1930
(Boissezon, 1930) that the nature of the fat reserves
accumulated in the insect body determined the maturation of eggs: sufficient reserves permitted ovary
development without a blood meal, whereas the blood
meal was required where fat reserves were insufficient. For Boissezon, the nutritional components influenced the reproductive patterns of Culex through a
purely physiological process, perhaps an endocrine
mechanism, as was later demonstrated (Boissezon,
1934). Roubaud rejected the proposition that nutritional behaviour determined racial distinctions
(Roubaud, 1934a,b). In the context of physiological
genetics, then developing principally in the USA and
Great Britain, races were indeed better associated
with genetic differences rather than being induced or
stabilized by the nature of food. It should be noted,
however, that the races of insects, particularly of
Culex, established by Roubaud have proven to be
valid and have long been used as genetically defined
variants (Clements, 1956).
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4
Social and economical perspectives
on Medical Entomology
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13 GILES ok:Layout 1
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Entomology in Translation: Interpreting French
medical entomological knowledge in colonial Mali
T. Giles-Vernick
Unité d’epidémiologie des maladies emergentes, Institut Pasteur, Paris.
Abstract. This essay examines how knowledge and practices around entomology and parasitology travelled and the consequences of their mobility. In exploring three anti-malaria campaigns in French Soudan
before 1960, it argues that the history of medical entomology’s travels entailed multiple temporal, spatial,
social translations that African medical personnel, intellectuals, healers, and farmers in French Soudan reinterpreted, appropriated, and sometimes wholly rejected. This essay also focuses on “erroneous” translations, detailing how and why middle class medical personnel and intellectuals interpreted and reformulated farmers’ and healers’ diagnostic categories that may or may not be malaria. Anti-mosquito and antilarval interventions, and more generally anti-malaria interventions, influenced how African colonial subjects and health workers understood certain vectors and of certain maladies. These understandings, in
turn, shaped the consequences of subsequent public health measures. Histories of translated parasitological and entomological knowledge and etiologies of illness have critical implications for contemporary
malaria control efforts: interventions to reduce malaria transmission through various kinds of entomological controls that require active participation of local populations cannot be effective if all participants cannot agree upon what is being controlled or prevented.
Key words: medical entomology, colonialism, Africa, ethnohistory.
“At every moment, translation is
as necessary as it is impossible”.
(Derrida [1999], 2004: 430)
This paper is about translation – and specifically
about how entomological knowledge about mosquitoes and malaria transmission was translated and
interpreted by diverse historical actors in French
Soudan, now Mali. From the early twentieth century, both parasitological journals and colonial
archives attest to the developing understandings of
the ecological, parasitological, and physiological
processes that Anopheline mosquitoes experienced
in transmitting malaria. But the translation of these
understandings in French Soudan, like all translations, was not simply about conveying ideas and
practices from one language into another (Vaughan,
1991; Fassin, 1993; Sadowsky, 1999; White, 2000;
Anderson, 2002; Anderson and Adams, 2007). In
the historical contexts of twentieth-century science,
French Soudan, and more recently, post-colonial
Mali, it also involved a transformation in ways of
evaluating the diagnostic categories, etiologies and
treatments of particular illnesses (Latour, 1999; Livingston, 2007)1.
Correspondence: Tamara Giles-Vernick, Unité d’épidémiologie des maladies emergentes, Institut Pasteur, 25 rue du Dr
Roux, 75724 Paris Cedex 15, France, e-mail: tamara.giles-vernick@pasteur.fr
1 Scholars (including most recently linguists, literary critics,
and anthropologists) have long and heatedly debated what
translation is and what makes a good one. Spivak urges the
translator to “surrender” to the text, to be “literal”. For Appiah, however, a literal translation is only a first step, and does
not necessarily convey the text’s meaning. What really counts,
he says, borrowing from Geertz, is a “thick” translation, one
Concerns about mobility, translation, and interpretation are at the heart of what Warwick Anderson and Vicanne Adams have called the “articulation of knowledge and practice across cultures”
(Anderson and Adams, 2007, 181; Anderson,
2002). Anderson and Adams have advocated studies that trace scientific knowledge production and
practice in multiple locations and that account for
the reasons and ways in which they travel (Anderson and Adams, 2007, 181-2). Such a vision can
help us to think about the mobility and reconstitution of entomological and parasitological knowledge
and practice.
This essay is a preliminary effort to explore one
small part of the complex voyages of medical and
entomological knowledge and practice. As many
studies of colonial medical practice in Africa reveal,
medical entomological knowledge did not simply
“diffuse” from France and then firmly implant itself
among populations in French Soudan (Anderson
and Adams, 2007; cf. Latour, 1999). Rather, colonial medical and public health experts translated
this knowledge to African medical personnel, who
re-interpreted it and re-translated it for farmers and
healers. These farmers and healers, for their part,
construed this knowledge in new ways, and they
sometimes rejected Anopheline mosquitoes’ role in
transmitting malaria. This essay argues that medical
entomology’s travels entailed multiple temporal,
spatial, social translations; it suggests that anti-mosthat relentlessly situates an expression in its cultural and historical context and conveys for learners (and not the author)
why such an expression is worth understanding. (Derrida
[1999], 2004: 427-8; Spivak [1992], 2004: 378, 379; Appiah
[1993], 2004: 394, 396-7, 400).
282
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T. Giles-Vernick - Interpreting French entomological knowledge in colonial Mali
quito and anti-larval interventions, and more generally anti-malaria interventions, influenced how
African colonial subjects and health workers understood of certain vectors and of certain maladies.
And these understandings, in turn, shaped the consequences of public health measures.
In tracing interpretations of three anti-malaria
campaigns before 1960, the essay also focuses on
“erroneous” translations, produced in the past and
at play in contemporary debates over malaria control in Mali. Over the past century, Malian middle
class civil servants, medical professionals, and intellectuals have embraced the insights of medical entomology, but they have simplified African farmers’
and healers’ knowledge as “mis-understandings” of
illness and its etiologies, attributing these groups’
limited acceptance of these biomedical insights to
several causes, including influential healers’ “charlatanism” and rural mistrust of western biomedicine.
In an analogous move, rural Bamana farmers and
healers have dismissed these middle-class interpretations as equally fallacious. “Erroneous” translations, however, should be of genuine interest to historians seeking to gain insight into the historical
misunderstandings and mistranslations that produced them, but also into contemporary malaria
control efforts: interventions to reduce malaria
transmission through various kinds of entomological
controls that require active participation of local
populations cannot be effective if ALL participants
cannot agree upon what is being controlled or prevented. As this analysis of translations and interpretations of three anti-malaria campaigns reveals,
there has never been such a consensus.
Sumaya, kónò, and the problems of
translating diagnostic categories
A conversation with a retired schoolteacher and a
medical student in the town of Niono provides a window into the historical accumulations of translated
entomological and medical knowledge. I had been
speaking to the teacher about the health consequences of a very large irrigated agriculture project,
the Office du Niger, in central Mali. The Office,
financed by the French and built by colonial subjects
in French Soudan from the 1920s, sought to reclaim
“desiccated” lands and to render them fruitful, irrigating cotton and rice fields with the waters of the
Niger River (van Beusekom, 2002, 8-11). I wanted to
know about sumaya – what researchers at the University of Bamako had told me was “malaria”. The
teacher explained that the Office du Niger had
brought water to a once-arid Niono, and with that
water came mosquitoes and sumaya. I then asked the
teacher about whether kónò (another disease whose
symptoms apparently resembled malaria) also
increased with irrigated agriculture. The teacher
expressed confusion, perhaps not understanding my
accent, and my research assistant, a medical student
named Makeou, attempted a translation into French:
“l’accès pernicieux”, he said.
L’accès pernicieux is a nineteenth-century term,
but in this context signified a very severe form of
malaria, transmitted by a mosquito of the Anopheles gambiae complex, which in turn infects a person
with falciparum malaria. What initially irritated me
– but then fascinated me – was how Makeou interrupted with a translation and how the teacher
immediately accepted it: kónò was without question
l’accès pernicieux. “Before,” she continued, turning
to me, “people attributed kónò to birds2. Here, it
was the owl that caused kónò, the owl that devours
children. When owls fly in the night, people were
afraid of them because they transmit the crisis.”
(Interview, F., Niono, 22 June 2006)3.
I came away from this conversation thinking that
in Mali, there were two diagnostic categories widely understood to be malaria: sumaya, an affliction
caused by the bite of a mosquito, producing fever,
chills, and headache; and kónò, a more severe illness afflicting children, who come under a malevolent force, usually associated with certain birds. My
assumption was that sumaya was a newer, medicalized knowledge, reflecting a century of western
entomological, parasitological influence. The historical puzzle, then, was to find out when, where,
through which networks and whom this entomological and parasitological influence had come.
But subsequent conversations muddied that onceclear correspondence between sumaya and kónò on
the one hand and malaria and mosquitoes on the
other. Mentioning these two disease categories,
sumaya and kónò, and their etiologies to a Malian
colleague, medical anthropologist Samba Diop, he
responded, “That’s not right. Those people know
nothing,” and he thus dismissed these etiologies and
categories of disease as faulty misunderstandings of
“local” medical and entomological knowledge. Continued conversations with healers (furakelaw) and
rural inhabitants of the Office du Niger revealed
that sumaya alone was not exclusively transmitted
by mosquitoes. Indeed, for many rural people with
whom I spoke, the importance of mosquitoes as
transmitters of malaria was dwarfed by other causes, from “unclean” foods to sugary ones. Some elaborated several other categories of illness that perhaps could be translated as malaria – but perhaps
not. For their part, middle class, western-educated
Malians insisted on reducing the complex etiologies
of rural Malians to mosquitoes and owls, and on
simplifying the disease taxonomy into two categories, sumaya and kónò.
These translations between rural Bamana-speaking Malians and middle class, educated Malian civil
servants have a history that neither straightforward
2
Doris Bonnet’s analysis of “la maladie de l’oiseau”, a diagnostic category widely shared among many societies in west
Africa, has been enormously useful (Bonnet 1999).
3
In compliance with the University of Minnesota Institutional Review Board, I do not divulge the identities of informants.
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biomedical translations of complex disease categories nor anthropologists’ symbolic interpretations
of complex local names and etiologies of illness can
capture. Straightforward biomedical translations
assume that there is a direct correspondence
between a disease like malaria and a host of indigenous diagnostic categories (Fegen et al., 2007;
Rieckmann, 2006).
But for Bamana speakers and other Malians (Jaffré 2003; Jaffré forthcoming; Diop, 2000; Diop,
2005; Roger, 1993), this has simply not been the
case. Indeed, anthropological translation of diagnostic categories illuminates causation from the perspectives of those who a disease afflicts (ScheperHughes, 1992; Whiteford, 1997; Olivier de Sardan,
1999, 7-12; Masquelier, 1999), but even these richly nuanced translations can run the risk of homogenizing and thus simplifying “local” understandings
of illness into the present. In distinction to biomedical translations, medical anthropological translations do “surrender” (Spivak [1992], 2004) to complex etiologies of illness (Olivier de Sardan, 1999:
7). The rich west African ethnographic literature on
concepts of sumaya, kónò, sayi, and other diagnostic categories evoking malaria grapples primarily
with present translations, and not with the historically sedimented nature of these disease categories.
Indeed, this literature, though scrupulously researched
and elegantly rendered, tends to depict such categories and their meanings as frozen in the present;
any evidence of change seems to indicate an “evolution” (Roger, 1993: 121), a term which only naturalizes the travels of biomedical knowledge as a
process of “diffusion”, without taking into account
the highly localized negotiations and frictions
through which scientific knowledge is both produced and interpreted (on “friction” see Tsing 2005;
Anderson and Adams, 2007; Livingston, 2007). In
distinction, while Jaffré’s work is not explicitly historical, it does offer possibilities for historical
insight into how and why people express within a
single category of illness multiple symptoms, etiologies, and indeed paradigms for understanding
human health (Jaffré, 1999, 161).
Historians have long shown that all medical
knowledge changes (Janzen, 1982; Feierman and
Janzen, 1992; Hunt, 1999; Livingston, 2007). The
multiple etiologies and symptoms of sumaya, reinterpreted through a historical lens, can reveal the
uneven ways that entomological and parasitological
knowledge traveled in the previous century. When
colonial health experts, scientists, African health
personnel, and colonial subjects translated entomological, parasitological knowledge into specific antimalaria practices, they engaged in conscious choices about the concepts, meanings, and practices that
they borrowed or rejected. The traces of these choices remain in the varied ways that people understand
and talk about illness. Moreover, translations took
place not only between participants in present clinical or public health settings (Tugwell, 2006), but
283
also across other kinds of social, economic and
political groups and over time. How should we
understand the contemporary use of a nineteenthcentury term, accès pernicieux, to refer to a set of
symptoms associated with paludisme? How have
the meanings and etiologies of sumaya changed
over the twentieth century? This essay thus offers a
preliminary historical analysis of translated diagnostic categories and etiologies, one that can allow us
to embrace the changing significance and the inconsistencies, complexities, and proliferations of specific categories, and to trace the peregrinations of
knowledge and practices around mosquitoes and
their control.
The travels of the mosquito:
entomological knowledge and practice
From the early twentieth century in Africa, antimalaria campaigns in effect put into practice –
indeed, translated – entomological and parasitological knowledge, seeking to prevent mosquitoes from
propagating and from coming into contact with vulnerable human populations (Opinel forthcoming).
In the context of West Africa, this “translation” of
medical entomological knowledge into public health
policy and practice requires further study. In the
colony that became French Soudan, such campaigns
took place throughout several circumscriptions over
the twentieth century. The earliest, in 1904, was a
campaign of “hygiène prophylactique” – a set of
environmental interventions to reduce mosquitoes
in European and African living spaces, and thus to
lower malaria transmission by limiting contact
between “fragile” European populations and infectious African ones, particularly children. “Hygienic
advice”, a 1904 pamphlet written by the Médecinchef LeMasle, contained documentation of diseasetransmitting mosquitoes, their habitats, and various
measures to avoid mosquito-borne and other diseases (LeMasle, 1904; Gouverneur Général de
l’Afrique Occidentale Française to Délégue permanent du Gouvernement Général, 1904; Gouverneur
Général de l’Afrique Occidentale Français to Monsieur le Délégué Permanent à Kayes, 1904).
LeMasle himself saw the goal of this pamphlet was
to translate “recent discoveries…on the important
role of mosquitoes in the propagation of certain
infections such as malaria” into prophylactic measures that rendered European life in Soudan healthier (LeMasle, 1904).
Although these reports and pamphlets were
intended to be read by French colonial officers, they
certainly shaped the living spaces of African colonial
subjects living in Bamako, Kayes, and other major
towns in French Soudan with substantial European
populations. Indeed, as in many other cities in colonial Africa in the early twentieth century, Bamako
became (at least on paper) a racially segregated city,
justified on the grounds that immunologically fragile Europeans required protection in the form of
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physical distance from African populations, and particularly African children (Curtin, 1985; Cell, 1986;
Watts, 1997, 261-63).
Subsequent efforts during the colonial period took
place in many circumscriptions of the colony, but
one well-publicized campaign occurred in the Office
du Niger, the massive irrigated development project
initiated by France in the 1920s and 1930s (van
Beusekom, 2002, xviii-xxix, 1-32; Echenberg and
Filipovitch, 1986). Tapping the waters of the Niger
River through a complex system of dams and irrigation channels, the Office irrigated arid lands north
of the Niger delta for rice and cotton production.
Under the French colonial administration and the
Service temporaire des irrigations du Niger (STIN),
African workers constructed a massive system of
irrigation, including the Markala barrage (begun in
1934, finished provisionally in 1941 and entirely in
1947) and an intricate network of canals and distribution channels to regulate the flow of waters. Originally envisioned to cover 1,850,000 hectares, the
irrigated area expanded only to 54,000 hectares by
1960 (van Beusekom, 2002, xxvii-xxix).
The Office du Niger 4, created in 1932, assumed
control of this irrigation system and in 1934 began
to recruit (often forcibly) African settlers to cultivate
its farms. After various sections of the irrigation system were completed, farming centers opened for cultivation and settlement between 1935 and 1952. The
region’s population thus grew rapidly, more than
doubling from 7,132 in 1935-36 to 14,861 in 193940 (van Beusekom, 2002, 157). Because of the long
process of construction, and in some locations, the
poor construction of canals and channels, farmers in
the Office long remained subject to seasonal changes
of the Niger River, suffering at times from flooding
and at others from dry conditions.
The Office’s irrigation system and agricultural
production scheme dramatically altered the once
arid lands beyond the Niger delta. Indeed, numerous
oral testimonies recount the ways in which the
Office irrigation system transformed the microclimate of the Niono region. As retired nurse H., one
of the very first Office employees to settle in Niono
in 1937, recounted
The Fala [river] was dry when I first arrived. [It had
been dry for] a very long time. You could see the
bones of fish at the bottom of the dry river bed.
There’s more rain now than there was before [the
Office]. The rainy season is now good. I arrived here
in June-July [of 1937]. In Bandiagara, July is really the
middle of the rainy season. But here in July, there was
only a little wind and dust at that time. I wondered
about this and asked someone, and I was told that the
wind and dust were the beginning of the rainy season.
The rains then were rare. If it rained two or three
times in a season, it was a good rainy season (Interview, H. Niono, 23 June 2006).
4 In this essay, the Office du Niger refers to the specific irrigation project, the region that it covered in French Soudan,
and the administration that oversaw this project.
In the mid-1930s, the newly-created Office provoked heated debates over whether irrigation had
precipitated an explosion of mosquito populations
and thus introduced or severely exacerbated malaria transmission. Another nurse vehemently insisted
that while the Office revived the “dead arms” (desiccated tributaries) of the river Niger, it also brought
huge populations of mosquitoes, and hence sumaya
– what he translated as malaria (Interview, F.,
Niono, 24 June 2006). Initially, the Office vehemently denied these claims, but eventually relented
and developed an extensive health care infrastructure, constructing a hospital at Segou, and several
clinics in its regions (Banguineda, Nienebale, Kokry,
and Niono), staffing them with auxiliary doctors,
midwives, nurses, as well as sanitary guards and
other workers. Nurses, performing the bulk of village health work, each assumed charge of three villages and paid daily visits to treat or evacuate the
sick and to distribute quinine as prophylaxis. Sanitary guards visited individual households, assuring
that Office farmers took steps to suppress the niches in which mosquitoes propagated. They further
ensured that no stagnant pools existed throughout
Office villages and that canals were cleared of vegetation and other debris. Yet available statistics indicate that the effects of these anti-mosquito measures
on malaria infection rates were mixed (Office du
Niger, Service Sanitaire, 1938; cf. Stapleton, 2000).
Anti-mosquito and malaria control efforts continued in French Soudan through the end of the 1940s,
focusing on household visits to identify and eradicate niches where mosquitoes propagated and to
distribute malaria treatment and prophylaxis (Service de Santé, 1949; Colonie du Soudan Français,
Service de Santé, 1950)5. But in 1950, the Service
de Santé also began applications of DDT in Bamako
homes, at least once a year in many locations but up
to four times during the year in those where
Anopheline populations were particularly dense
(Colonie du Soudan Français, Service de Santé,
1950) 6. Available documents through 1954 indicate
that DDT spraying continued in Bamako, but it
remains unclear precisely when it expanded into
other regions of French Soudan. By 1957, however,
the Service d’Hygiène Mobile had expanded into
five zones, including one containing the Office du
Niger (Colonie du Soudan Français, 1957).
Interpreting anti-mosquito and anti-larval practices
This essay does not address the measurable morbidity and mortality effects of these campaigns (cf.
Moulin, 1996), partly because the data are highly
5 On research relating to malaria treatments, see Sweeney,
2000. Drugs used for treatment and prophylaxis included quinine, premaline, and nivaquine.
6 This campaign had followed DDT trials elsewhere and
had preceded the WHO’s Global Malaria Eradication Programme in 1955-56 (Litsios, 2000).
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inconsistent and thus treacherous to interpret, but
primarily because I am more interested in how
practices around mosquito control (themselves
translations of research into entomology and parasitology) shaped how people thought about the diseases that afflicted them. Because this analysis is
based upon historical field research and published
ethnographic accounts, it remains treacherous to
link particular recollections or perspectives on
insects and fevers to specific campaigns over the
twentieth century. Rather, I interpret the oral historical accounts as expressions that simultaneously
engage the past and present (Tonkin, 1992), that
contain sedimented understandings of past changes
and influences. Such contemporary accounts incorporate evidence of past (but undateable) influences
(Giles-Vernick, 2002). Moreover, the evaluations
here come from informants in the Office du Niger
and in Bamako, who have accumulated diverse
understandings of mosquitoes and their roles in
transmitting disease.
The influences of these malaria campaigns on
middle class concepts of sumaya — which they
translated as paludisme or malaria – were evident.
For urban civil servants and intellectuals in Bamako
and the Office, as well as for medical personnel
who had worked in various health services and on
anti-malaria campaigns, sumaya was a medicalized
category of disease, transmitted exclusively by mosquitoes (undifferentiated moustiques) and characterized by fever, chills, and headache. For these particular groups, the travels of this knowledge took
similar routes: many of the middle class intellectuals, civil servants, and medical personnel interviewed for this study were the children and grandchildren of African colonial subjects who lived in
the urban centers of Bamako, Kayes, Sikasso or in
the Office. They either interacted with colonial
anti-mosquito and anti-malaria interventions as
nurses, schoolteachers, clerks, and other employees
of the colonial administration, or they were the
children of colonial personnel who had done so (M.
and M., Niono, 23 June 2006). H. and M. for
instance, spoke of coming to work as nurses for the
Office du Niger in the 1930s and 40s; F. arrived in
Niono as a nurse in the late 1950s during the DDT
campaigns, to work as a nurse for the Office du
Niger; N., however, trained as a nurse but worked
as a technician in the 1950s, conducting studies on
French Soudan’s mosquito species and their transmission capacities (Interviews: H., Niono, 23 June
2006; M. 20 June 2006; F., Niono, 24 June 2006;
N., Niono, 23 June 2006).
These informants’ understandings of the malaria’s etiologies and effects appear to have shaped
their interpretations of subsequent malaria interventions. Many, for instance, lauded the DDT
spraying campaigns in the 1950s and 60s and
embraced current efforts to avoid contact with
mosquitoes by using impregnated bednets (Interviews: B., Bamako 30 June 2006; N., Niono 24
285
June 2006; O., Bamako, 10 June 2006; T. Niono,
23 June 2006)7.
Moreover, their altered understandings of the disease’s etiology also affected their perceptions of rural Bamana populations’ past and present concepts of
illness. Indeed, as explained earlier, many erroneously represented rural farmers’ etiologies of
paludisme, invoking categories of sumaya and
kónò. While rural people distinguished these two
categories as having different etiologies and symptoms, educated Malians in the Office du Niger and
Bamako contended that they were the same illness –
malaria. Middle class depictions of rural Bamana
knowledge suggested that undue confidence in
arcane (and in their perception, exotic) treatments,
mistrust of white colonizers’ medicine, and a lack of
education explained the persistence of alternative
etiologies. Malian schoolteacher M., explained,
Peasants took care of children with kónò by calling
healers. The healers would massage the child and say
incantations. And if the child urinated or defecated,
he was saved. But if not, the child would die. It was
old people – men and women – who were healers.
They didn’t have any medicines, but would use incantations.
TGV: Why was urinating a sign that the child would
live?
M.D. [M.’s husband]: With kónò, the child’s body is
completely stiff. If a child could allow fluid to escape
his body, it meant that there was a sign of life in the
body.
M. People thought that when a child urinated, that
meant that the owl had left the body of the child.
TGV: And so when did people begin to embrace western medical concepts of malaria?
M.D.: Before, people weren’t educated. Very few had
the advantage of going to school….Most people had a
lot of confidence in medical tradition. Their religion
was mixed into it. If you went to a doctor to get an
injection, it was a fearsome experience. But people
did know how to pick a plant, boil it, and drink it.
People were afraid [of doctors and their medicines].
There are people around who are over 80 years old
and have never had an injection, never taken a pill.
Now, there are more intellectuals, more hospitals, and
people have begun to have confidence in modern
medicine. During the colonial period, there were very
few doctors in Mali, and the Malians who were
trained weren’t even doctors: they were only infirmiers d’etat. (Interview, M. and M.D., Niono, 23 June
2006).
Although these teachers suggested that rural beliefs
around the causes of sumaya and kónò were changing, other middle class health care workers, teachers, and other civil servants insisted that these older etiologies still held sway with rural farmers and
healers (Interviews: F., Niono, 22 June 2006; F.,
Niono, 24 June 2006; N., Niono, 24 June 2006; T.,
Niono, 24 June 2006; O., Bamako, 2 July 2006; S.,
7
I asked relatively little about middle class, educated informants’ understandings of Mali’s anti-malaria efforts but
intend to pursue this dimension of the study in the future.
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3 July 2006; B., Bamako, 19 June 2006). Indeed,
one schoolteacher insisted that farmers continued to
consult “charlatans-guerisseurs” (“charlatan-healers”), whose botanical remedies were only sometimes efficacious but whose incantations were “useless”.
Such perceptions, as scholars have noted elsewhere, have their historical roots in colonialism, as
medical missionaries and colonial health professionals offered western medicine as a means of dispelling subjects’ ignorance, of civilizing these colonial subjects, and of incorporating them into colonial projects (Pigg, 1997, 262). While Stacy Leigh
Pigg notes that Indian nationalists embraced “traditional medicine” to assert their autonomy, (Pigg,
1997, 262) it seems that in French Soudan and contemporary Mali, at least middle class teachers,
researchers, and medical took up biomedical conceptions of Anopheline mosquitoes as transmitters
of malaria and dismissed some rural healers’
(furakelaw) knowledge and practice, perhaps as
markers of their own “modernity” forged in educational institutions, and of social differences between
them and farmers. It is also worth noting that perhaps middle class informants had not appropriated
this etiology as completely as they claimed; two
healers asserted that civil servants sought out their
skills frequently to heal their children (Interview,
M., Niono, 25 June 2006; M.R., 26 June 2006). In
2007, I watched an internationally recognized
Malian malariologist diagnose his son with malaria,
give him an artemisinine combination therapy, and
call in a trusted healer to treat the boy.
Additional field research with farmers and healers
themselves yielded a substantially more tortuous set
of etiologies and symptoms of diseases that may or
may not be malaria. In this case, too, successive
anti-malaria campaigns have influenced these populations. Samba Diop, Yannick Jaffré, and others,
have worked for many years with Bamana healers,
authoring several pieces examining their contemporary ethno-medical knowledge (Diop, 2000; Diop,
2005; Jaffré, 1999; Jaffré, 2003, forthcoming; Roger
1993). In one study exploring access to malaria
treatments in several Malian villages (Diop, 2000),
Diop painstakingly elucidates at least six complex,
sometimes overlapping illnesses that can evoke or
designate malaria, all with diverse etiologies ranging
from the physical effects of mosquitoes, to sweet,
cold, or fatty foods, to wind and humidity, to the
malevolent forces of birds and chameleons (see also
Roger, 1993). Indeed, these elements can throw
individual human bodies into a kind of disequilibrium, but certain bitter medicinal plants can re-equilibrate that imbalance. Of sumaya itself, he reports
that his informants also perceived it as a condition
that was an integral part of one’s blood: “One can
never be without sumaya; it is indistinguishable
from the blood of living people” (Diop, 2000, 37).
These concepts were echoed in oral histories with
elderly healers and farmers in Niono about the
health effects of Office irrigated farming and antilarval campaigns. These diagnostic categories lend
themselves to a more historical interpretation. Take,
for instance, the category sumaya, which appears to
have accumulated a series of historically produced
meanings. One farmer, who contended that the
Office du Niger introduced malaria to the region,
argued that the term sumaya had once referred to
an entirely different illness, one contracted during
the region’s very lengthy dry season, and with entirely different symptoms. It was only later, he argued,
with the arrival of Malian nurses and French medical officers that the term sumaya came to mean
paludisme, or malaria. And indeed, early twentieth
century scientists in French Soudan mentioned a
disease, “soumaya” or “souma”, but translated it as
bovine or equine trypanosomiasis (Cazalbou, 1905,
564; Bouffard, 1907, 71-3). These claims all merit
additional investigation: what was sumaya or
souma for early twentieth century Bamana speakers? Why did French scientists translate it as bovine
or equine trypanosomiasis? Why did its significance
change, for whom, and when?
Among rural inhabitants in the Office du Niger,
sumaya’s diverse causes should thus be interpreted
not only as contemporary evidence that an illness
can be provoked by multiple causes, but also as evidence of circulating knowledge, practices, and
objects associated with colonialism (Hunt, 1999).
French entomological and parasitological knowledge
concerning Anopheline mosquitoes and malaria
transmission, translated into practice through antimosquito campaigns, was only one such influence,
and it clearly has never stabilized in some rural
Bamana speakers’ etiologies of illness. Indeed, M.
Roger found over a decade ago among women informants in Sikasso (further south in Mali) that precisely what mosquitoes did to transmit sumaya was
widely debated: mosquitoes injected “dirty water”,
the blood of other people, its own blood, or some
kind of contagious agent that caused people to grow
ill (Roger, 1993, 98).
Moreover, mosquitoes as vectors existed alongside
other etiologies (an insight that Roger also points
out, but relegates it to “incomplete understandings”
of mosquitoes as vectors). Mama R., an elderly healer or furakela, well known in the Office region and
beyond for her work with children, for instance,
contended that sumaya was caused by “water that is
not clean, foods that are not clean…or sweet foods”
(Interview, Mama R., Niono, 25 June 2006; Jaffré,
1999). M.C., another woman, contended that while
mosquitoes could provoke sumaya, fresh unboiled
milk and unclean foods like rice and millet could as
well (Interview, M.C., Niono, 25 June 2006). And
another well known healer whose clients sought her
out from various regions of West Africa concurred:
Many people say that it is caused by mosquitoes. But
it isn’t just that. It also has to do with the age of the
child and how the parents of child feed that child.
Some foods have impurities in them (dumuni
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nogolen). Those impurities stay in the body and can
provoke sumaya (Interview, M.D., Niono, 26 June
2006).
Healers or farmers also held starkly different understandings about the relationship between sumaya
and kónò, some distinguishing it on the basis of etiology, others on symptomology, and still others on
the treatment. Mama R., for instance, staunchly
asserted that
Sumaya and kónò are not the same. Kónò is caught
by children who have a kind of crisis. They are dry,
feverish. To treat kónò, you take leaves of ntonge,
which is a tree. You cut the leaves, boil them in water,
and give them to the child to drink. When the child
urinates or defecates, then he is cured.
When I asked how kónò was provoked, she answered,
When a pregnant woman sees an animal, some kind
of animal, and throws a stone at it and hurts it, then
the child she carries can get kónò. When you’re pregnant, you can’t hurt another being. It can also be
caused by birds – but lots of different kinds of birds
– chickens, guinea fowl….After the woman gives birth
to the child, the child can get it. (Interview, Mama R.,
Niono, 25 June 2006).
A.S., the wife of an Office du Niger farmer, seemed
to hedge her bets with me when I asked her about
sumaya and kónò, claiming that she took her children to the dispensary when they had sumaya, and
that “people at the hospital say that [kónò] is malaria, and farmers who work for the Office think it is
malaria” (Interview, A., Niono, 24 June 2006). In
the villages, she continued, many people disagreed
and would massage a child with kónò until the child
urinated or defecated, indicating a full cure. As for
another Niono farmer, S.S., who worked for the
Office, he asserted, “Some people say that kónò is
sumaya, but it isn’t. It’s a different illness. It’s
caught by children. Their eyes open wide, and they
become very dry”. Yet another healer, M., insisted
that sumaya and kónò “are sort of the same thing,
but there is a little difference between them. When
you have had sumaya for a long time, it can turn
into kónò. The child’s eyes become yellow and the
child vomits”. (Interview, M., Niono, 25 June 2006).
She elaborated other conditions loosely related to
sumaya – sayi – a diagnostic category widely shared
among Manding peoples, primarily characterized by
yellowness (of eyes, of vomit, of urine), and explored
in detail by Y. Jaffré (Jaffré, 1999, 155, 161).
It is not at all surprising that informants simultaneously embraced diverse etiologies of these maladies. Medical pluralism in Africa and elsewhere has
been explained in divergent terms: as “intentional
hybridity” (“Inherently political, a clash of languages
which question the existing social order”) (Werbner,
2001, quoted in Marsland, 2007, 755); but also as
“productive misunderstanding” (Livingston, 2007;
see also Livingston, 2005, and Fassin, 2007). My
research suggests that both models may be appropriate, and that these diverse understandings of sumaya
and kónò should be read historically, as the sedi-
287
mentation of older and more recent concepts that
have linked particular health consequences to certain
environmental exposures (foods and birds, for
instance). For urban and middle class intellectuals,
who have appropriated biomedical diagnostic categories and treatments, sumaya’s possibly older etiologies and symptoms appear to have far less currency; older notions of sumaya simultaneously allow
educated middle class people to hedge their bets, but
also to distinguish themselves from rural people who
have not fully accepted modern biomedicine. But
additional investigation is needed. African participants – the infirmiers, auxiliary doctors, sanitary
guards, and inhabitants of locales in these past campaigns – were well placed to translate and to integrate these efforts into their own understandings of
fevers (and perhaps malaria).
For rural farmers and healers, the coexistence of
sumaya’s multiple etiologies (mosquitoes who transmit a range of infecting substances, sweet foods,
impurities) can be interpreted as a selective, uneven
appropriation of colonial medical and entomological
knowledge that was translated into successive antimosquito and anti-larval campaigns in colonial and
contemporary Mali. But following Livingston, the
consequences of this piecemeal appropriation has
been that biomedical paludisme on one hand and
sumaya, sayi, and kónò on the other, appear to
remain distinct. For these populations, cold (refrigerated) foods and certain understandings of cleanliness and filth (dirty water and foods) may also have
constituted colonial markers of modernity, but ones
that made people sick. Could this etiological plurality reflect their historical ambivalence of this modernity? This question, too, merits further investigation. Moreover, these colonial concerns, still evident
in present concepts of fevers, may coexist alongside
older concerns about how sweet foods interact with
human bodies. It also appears that a much older etiology and symptomology of sumaya as a dry season
affliction with different symptoms has disappeared
almost entirely.
Nevertheless, it is clear that French entomological,
parasitological knowledge around malaria, translated through anti-malaria campaigns, was less actively appropriated by rural Office farmers and healers.
As both rural farmers and middle class intellectuals
and health care workers contend, rural people have
historically had diminished access to biomedicine
because of poverty, long distances from health care
infrastructures, but also because of their substantial
mistrust of biomedicine until at least independence
(Interviews: S.S., Niono, 24 June 2006; A.S., Niono,
24 June 2006; F.D., Niono, 24 June 2006).
Conclusion
In 1999, the anthropologist J.-P. Olivier de Sardan
argued, “Most popular representations of illness
[in Africa] which are still in circulation were produced in precolonial languages and cultures, with-
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out any relationship with modern medicine. Of
course, in contact with this [modern medicine]
since colonisation, they have evolved, and new representations have been added to old ones. But the
daily relationships that people today maintain with
illnesses continue most often to be thought of and
spoken of with the words and categories little
influenced by western biomedicine, and which are
a part of the ‘basis of local culture’” (Olivier de
Sardan, 1999, 7).
This essay, while indebted to the rich contributions made by west Africanist medical anthropologists, began with a historian’s assumption: Malian
medical knowledge, practice, and diagnostic categories have changed. The aim here has been to
explore how the travels of medical entomological
knowledge and practice, translated into certain
twentieth-century public health measures, influenced Malian understandings of scientific entomological and parasitological knowledge and practice
around malaria. Anti-mosquito and anti-larval
efforts, conceived by colonial public health officials,
were literally translated into Bamana language and
practice through African doctors, nurses, and sanitary guards to varied colonized populations in
Bamako and the Office du Niger. Exploring these
multi-layered translations of entomological knowledge and practice across languages (French and
Bamana), but also over time (early 20th century to
early 21st century) and across social groups (middle
class health personnel and intellectuals and rural
populations) not only help to excavate the social
and cultural understandings of certain illnesses
among Bamana speakers. They can also illuminate
the complex, sedimented, and uneven influences of
itinerant medical knowledge and practice, in this
case the ways that entomological and parasitological
knowledge about Anopheline mosquitoes and
malaria transmission has in some cases displaced,
but also coexisted with etiologies of fevers that may
– or may not be – malaria 8.
Additionally, this essay has suggested that an
analysis of “erroneous” translations, past and present, may be illuminating. While Malian middle
class civil servants, medical professionals, and intellectuals have characterized African farmers’ and
healers’ conceptions of “malaria” as “mis-understandings” of the illness and its etiologies, rural
Bamana farmers and healers and intellectuals sympathizing with them have dismissed these middleclass interpretations as equally fallacious. These distinctions in disease taxonomy and etiologies appear
8
These insights thus reframe Randall Packard’s important
question, “what is malaria?” (Packard, 1997). Malaria, he
observes, has been understood narrowly as illness caused by
a parasite that is spread by mosquitoes, but also at times as
“a problem of social uplift and thus ultimately tied to social
and economic conditions, like rural housing, nutrition, and
agricultural production” or the result of failed economic policies”. But the present analysis asks: What is not malaria? For
whom? And when?
to have been produced historically, the consequence
of different African populations’ engagements with
successive anti-mosquito, anti-larval, and antimalaria interventions.
Malian nurses, teachers, and scientists consolidate
multiple and overlapping disease categories of rural
farmers and healers, but they also recognize, albeit
in a limited way, the complexity of Bamana beliefs
in articulating differences between sumaya and
kónò. But what is at stake for Malian intellectuals in
this translation? By embracing a biomedical etiology of malaria and representing a reductionist understanding of rural Malians, how do they bolster their
own positions and represent rural Malians? It is
clear that Malian intellectuals have come to occupy
positions as intermediaries between rural populations and international donors and organizations.
What remains to be studied are the stakes involved
in their simultaneous homogenizing and representing complexity.
While erroneous translations can illuminate the
historical and political investments in these translations, it is worth keeping in mind that they have their
shortcomings, and that malaria control efforts need
to aim in the present and future for a better understanding of the diverse categories and etiologies of
illness. Perhaps public health agents within and outside of Africa have realized limited successes in their
efforts to control malaria because they have persistently reduced complex etiological factors and several categories of illness to a single “malaria”. But the
stakes of how we translate are high (Farmer 1999
and 2003; see also Charles Rosenberg’s discussion of
a “shared understanding” of illness in Rosenberg,
1992, and Rosenberg and Golden, 1992), for malaria continues to extract a considerable human toll in
Africa (Sambo, 2007, iii).
Many African states, including Mali, have sought
to address this toll. In 1993, for instance, Mali
launched its Programme nationale de la lutte contre
le paludisme (PNLP), and over the past several
decades, it has developed a series of long-term plans
to reduce mortality and morbidity from malaria
(Maïga, 1994; Konate, 1993, 84, 86-7). Mali has,
moreover, undertaken these considerable efforts in
collaboration with bi-lateral and multi-lateral organizations (World Bank, the World Health Organization), philanthropic foundations, and private corporations, which have provided substantial financial
and material resources for research and anti-malaria campaigns (Multilateral Initiative on Malaria,
Roll Back Malaria Partnership) (Glass and Fauci,
2007: iv-v; Rugemalila, 2007, 296-7; Konate, 1993,
10-11, 14; for an evaluation and critique of Roll
Back Malaria, see Packard, 2007, 220-246).
Randall Packard has recently argued that interventions to control malaria must account for the
fact that “[m]alaria is a multisectoral problem that
needs to be attacked on multiple fronts…”, and that
such efforts must be sustainable (Packard, 2007,
248-250). I would add to Packard’s compelling
26-08-2009
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Pagina 289
arguments that contemporary public health interventions into malaria have not truly sought to grapple in any fundamental way with diverse understandings and etiologies of illness, and to incorporate them into their interventions. Most frequently,
these diverse categories are effectively collapsed into
a single malaria and are accompanied by calls for
more public health education. But “more public
health education” is premised on a faulty assumption that people will change their conceptions of
malaria and other illnesses, jettisoning older concepts of illness, once they gain access to that education. And yet parasitological and entomological
knowledge have circulated (albeit unevenly) in
French Sudan and independent Mali for more than
a century – and older understandings of illness that
may have some relationship with malaria remain.
The sustainable interventions that Packard calls for
are not only economic and social, but also intellectual. Indeed, public health institutions need to begin
their efforts by building on diverse meanings and
causes of illness, or their best efforts will be wasted.
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Science and popular participation in the investigation of
heartwater in South Africa, c. 1870-1950
D. Gilfoyle
The National Archives, Kew, London, UK.
Abstract. During the late nineteenth century, settler farmers in southern Africa identified heartwater as a
damaging disease of small stock and cattle. They advanced various explanations of the disease, including the theory that it was caused by the bite of ticks. Around 1900, the American entomologist C.P. Lousbury demonstrated that heartwater was transmitted by the bont tick. He also worked out the life cycle and
life habits of the tick. Subsequently, farmers developed methods of controlling ticks by dipping animals
in solutions of arsenic. By 1910, the practice of dipping cattle had become very widespread over much
of southern Africa. The expansion of the practice was greatly stimulated by the coming of the deadly tickborne disease, East Coast fever. At this time, veterinary scientists attempted to develop a vaccine against
heartwater, but with little success. Little further progress was made until the 1920s, when the American
scientist E.V. Cowdry identified a causal agent, Rickettsia ruminantium, while on a research secondment
to South Africa. By the 1940s, South African veterinary scientists had devised methods of immunising
stock against heartwater, but there remained considerable technical difficulties and their use remained
limited. Dipping in arsenic solutions to attack the tick on the animal thus remained the most important
means of controlling disease in the first half of the twentieth century.
Key words: heartwater, tick-borne diseases, veterinary medicine, South Africa.
Tick-borne stock diseases, such as East Coast fever,
heartwater, babebiosis and anaplasmosis have historically shaped pastoralism in southern Africa. The
control of these diseases has been an important element of state veterinary policy and has entailed the
deployment of considerable resources. Of these diseases, only East Coast fever has received significant
attention from historians, who have examined scientific research and political opposition to government policy in the first twenty years of the twentieth century (Cranefield, 1991; Bundy, 1987; Giblin,
1990; Waller and Homewood, 1997).
This article examines the historical encounters of
scientists and colonial farmers with heartwater
between the 1870s and 1950. During the 1870s,
colonial scientists depended heavily upon the white
farmers with whom they predominantly dealt for
information about the disease. I examine the rise of
laboratory studies of heartwater in the twentieth
century, but argue that farmers continued to play a
role in the construction of a body of knowledge
about heartwater through their participation in field
experiments. I am concerned with the practices of
veterinary scientists and entomologists, as much as
with knowledge and theory (Worboys, 2002) . In a
society in which European farmers held considerable political power, this knowledge was constituted
mutually by colonial scientists and the settler farmers with whom they primarily dealt. Accordingly, I
emphasise the importance of popular ideas and
Correspondence: Daniel Gilfoyle, D. Phil., The National
Archives, Kew, London, UK; Research Affiliate, Wellcome Unit
for History of Medicine, University of Oxford, UK, e-mail:
daniel.gilfoyle@nationalarchives.gov.uk
practices in shaping veterinary knowledge and preventive technology.
I also contextualise the material within some wider
themes in the history of colonial veterinary and
medical science. Michael Worboys has argued that
following the “stamping out” of the 1865 rinderpest
epizootic in Britain, germ theory was quickly accepted within the British veterinary science, although
the consequence was the entrenchment of the regulatory regime rather than the adoption of germ practices, such as preventive inoculation (Worboys,
2002). Until the 1930s, the large majority of vets
working in South Africa had trained in Britain, but
the trajectory of veterinary science and policy was
quite different. I examine how veterinary scientists
in South Africa responded to the particularities of
the locale by devising strategies of control based on
the control of ticks and the use of prophylactic inoculation. This article is also concerned with the interactions of veterinary scientists in southern Africa
with practitioners and institutions in other parts of
the world. David Wade Chambers and Richard
Gillespie have recently challenged a tendency in the
historiography to interpret colonial science as essentially oriented towards and communicating with an
imperial, metropolitan centre. Instead, they describe
modern science as a “polycentric communications
network” of which colonial science formed a part
(Wade Chambers and Gillespie, 2000). This article
uses research into heartwater as a case study in
which to illuminate some international scientific
interactions.
The first section of this article discusses some veterinary and popular ideas about the cause of heart-
292
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D. Gilfoyle - The investigation of heartwater in South Africa
water between 1870 and 1902, concentrating on the
discovery of the tick vector. It discusses the impact
of this discovery on methods and technologies of
disease control. The second section considers microbiological studies on heartwater during the 1920s.
Connections are drawn with research in the United
States, but it is argued that gains in knowledge
about the cause of the disease had relatively little
impact upon methods of control. These continued to
concentrate on attacking the tick vector on the animal. The final section examines experiments with
prophylactic inoculation. To the 1940s, farmers continued to play role in the generation of knowledge
about disease.
Ticks and disease:
some popular and veterinary ideas
In 1876, the Cape government responded to an
apparent proliferation of stock disease, which was
damaging the pastoral economy, by appointing a
Colonial Veterinary Surgeon and setting up a Stock
Diseases Commission (Beinart, 1997). The Commission identified scab and internal parasites as
important causes of loss, but in the wool-producing
Eastern Cape districts, more obscure diseases were
reported to be common (Gilfoyle, 2002).
In giving evidence to the Commission, John Webb,
who farmed in Albany district linked an upsurge in
disease to the introduction of the bont (variegated
or tortoiseshell) tick. This tick, Webb alleged, had
first been noticed in Albany on cattle brought from
Zululand during the 1930s. Since then it had proliferated and spread across the district. Webb
thought that fatal diseases, “gallsickness” and
“boschsickness” (bushsickness) in cattle and “heartwater” in sheep, were caused by “inflammation
brought on by the tick” (CPP, 1877, 108-9 and
135). One Fort Beaufort farmer, Bezuidenhout, testified that the bont tick had first been observed
around the Gonubie River in 1835 and had gradually invaded surrounding districts, spreading disease
wherever it appeared (CPP, 1877, 129). Other farmers described how the bont infested bushy places
only, avoiding the open veld; sheep and goats which
remained healthy on the open ridges died of heartwater if taken into the bushy ravines (CPP, 1877,
115 and 120).
Albany farmers were familiar with ticks, which they
considered to be generally harmful, and one identified four distinct types: the bont, blue, skilpad (tortoise) and the small red ticks (CPP, 1877, 118). The
bite of the large bont tick, it was said, could destroy
the nipples of a milch cow, while smaller specimens
caused lameness by lodging between the “klauws” of
a sheep’s foot. The small red ticks seemed to cause
paralysis in lambs, which might die if the ticks were
not removed. But they did not necessarily agree that
ticks caused “heartwater” or “gallsickness”, the
obscure diseases which, it was claimed, were ruin-
ing wool production in the Cape”s eastern districts
(CPP, 1877, 115 and 118). Some argued that overstocking, with the consequent decline in the quality
of pasture, was the cause of the new diseases (CPP,
1883, 36). Hobson, a Jansenville farmer, linked outbreaks of heartwater in the interior to ox-wagon
transport from coastal districts such as Uitenhage.
He believed that pastures on which transport oxen
from the coast were allowed to graze would subsequently become fatal for goats (CPP, 1883).
Some of these arguments were taken up by farmers
in the pages of the Cape”s Agricultual Journal, first
published in 1888. Andrew Smith, an Albany
farmer, published an article in 1889, which sought
to account for the virtual disappearance of sheep
farming from districts such as Victoria East, Lower
Albany and East London, which had once held some
of the best sheep runs in the colony. Smith argued
that the immediate cause of the decline was a deterioration of the veld caused by overstocking. Sheep
ate out the nutritious sweet grasses first, leaving the
poor grasses and allowing opportunistic noxious
weeds, such as ragwort, to gain a hold. (For ideas
and policies on poisonous weeds, see van Sittart,
2000). The plants which Africans used as “germkillers and anti-septics” were among the first to go.
The soil became depleted of mineral salts (potash,
lime and phosphorus) so that the “balance of replenishment is overturned” and animals starved as a
result. Disease had also played a part in the decline
because the proliferation of sheep in the early years
of farming had poisoned the pasture. “Diseased
sheep pass out the bacterial spores and the eggs of
low forms of animal life”, which, Smith claimed,
were able to survive for a time on the veld to infect
susceptible animals (AJCGH, 1889). Smith’s article
thus encapsulated and combined a number of popular theories about the causes of stock disease:
nutritional deficiency; plant poisoning and a version
of germ theory which was linked to “worm theory”.
Smith’s ideas stimulated a debate in the correspondence pages of the Agricultural Journal over the
next few years. William Rogers, who had moved
from Albany because of the prevalence of stock disease there, inverted Smith’s argument about overstocking. Rogers claimed that the decline in sheep
farming in these Eastern Cape districts was primarily due to diseases, identifiable by the accumulation
of fluid around the heart and swollen gall bladder,
which were known as “heartwater” and “gallsickness”. These diseases had spread, since their first
appearance around 1860, slowly up the valleys of
the Eastern Cape and across contiguous low-lying
land. Rogers was impressed by the localised occurrence of disease and its apparent correlation with
altitude; like witnesses to the Stock Diseases Commission, he noted that sheep still did well on higher ground in “heartwater” districts, although nearby
kloofs and valleys, characterised by a more luxuriant vegetation, were fatal. Overstocking had,
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through constant manuring, actually enriched pastures to the extent that they became poisonous,
especially in the “semi-tropical” environment of the
valleys (AJCGH, 1890a). L.I.R., an Adelaide farmer,
speculated that germs which were unable to survive
on “sour” veld, might be able to live on veld which
had become “richer and sweeter” (AJCGH, 1890b,
86).
One Sandflats farmer agreed that overstocking
could lead to veld deterioration, but argued that in
the case of the Eastern Cape the decline of sheep
farming was caused specifically by heartwater.
Heartwater was a “germ” disease, but one that
depended on certain vegetation types for its propagation. The difference in vegetation between, for
example, the valleys of Victoria East and the higher
ground of Bedford accounted for the presence of the
disease in the former district and its absence from
the latter (AJCGH, 1890c, 45). In particular, he
linked the incidence of “heartwater” to the presence
of thorny mimosa or acacia (AJCGH, 1891, 124).
A Fort Beaufort farmer, Ralph, advanced what was
perhaps the opinion of “the majority of the more
experienced farmers” in reiterating John Webb’s evidence to the Stock Diseases Commission. The disease was caused by the bont tick, which, Ralph
claimed, was never absent when the disease
occurred. The adult female, “about the size of a
medium plum”, was capable of producing thousands
of eggs and posed, he argued, a worse threat to the
Colony than scab. Legislation similar to the Scab
Act, and the appointment of inspectors would be
necessary if the tick was to be eradicated (AJCGH,
1890d, 68). Farmers did not, however, perceive
ticks as transmitters of germs. These comparatively
large and highly visible creatures were described as
a cause of disease in themselves, and something
with which farmers could perhaps deal.
Duncan Hutcheon, who was appointed as the Cape’s
sole government veterinary surgeon in 1878, examined many post-mortem cases of heartwater. They
revealed a coagulating effusion of a “pale strawcoloured fluid” in the heart sac, suggested to him that
this was distinct from the common “dropsy” of worm
infestation also referred to by farmers as “heartwater”
(CPP, 1882a, 6; Beinart, 1997, 240). With the recent
work of Koch and Pasteur on anthrax in mind,
Hutcheon thought heartwater was a “specific” disease
caused by a spore-forming bacillus, which contaminated the veld (CPP, 1882b, 1). Apart from what was
already common knowledge, however, he had little to
offer in the way of advice.
In 1892, Drs Smith and Kilborne of the United
States Bureau of Animal Industry published a study
showing that Texas fever, a protozoan disease of cattle, was transmitted by ticks (Smith and Kilborne,
1893). An article in the Agricultural Journal
described these findings in some detail and speculated that Texas fever was “a plague probably not
293
differing specifically from our colonial Redwater”
(AJCGH, 1992, 202; Beinart, 1997, 247). Gradually, the possibility of tick transmission took hold
among the small body of vets working at the Cape.
In 1897, Robert Koch, who was researching a
rinderpest vaccine in Bechuanaland, saw a parasite
identical to that of Texas fever in blood samples
(CPP, 1897, 9). By 1899, the vets were taking the
question of tick infection very seriously.
The tick transmission of heartwater was worked out
by the Cape’s government entomologist, Charles
Lounsbury, an American scientist who had been
employed at the Cape since 1895, initially to investigate pests of fruit (Brown, 2003). He argued that
the beliefs of many farmers deserved investigation
and an effective preventative might re-open the
south-eastern districts of the Colony for sheep farming and wool production (Lounsbury, 1899, 729-31;
AJCGH, 1901, 305). He contacted Hutcheon, suggesting co-operation between the veterinary and
entomological branches of the Agricultural Department (CAD, 1898).
Lounsbury began his research on the bont tick late in
1898 using facilities provided by the Fort Beaufort
dairy farmer, Llewellyn Roberts, at the Cottesbrook
creamery, Adelaide (Lounsbury, 1899, 742). He identified the bont tick as Amblyomma hebraeum, which
had been classified by the German scientist C.L. Koch
50 years previously. The eggs of the bont failed to
hatch unless they were kept continually moist and in
the shade, suggesting that its range would probably be
limited to the wetter, more densely vegetated regions
of the Cape and confirming the farmers’ correlation
of the disease with damp, bosky habitats. The bont
tick remained on its host for only about eight days at
each stage of its life cycle (larvae, nymph and imago),
living on the ground during the intermediate periods.
The blue tick, which transmitted redwater fever, was
a more sedentary creature, spending most of its life on
the animal (CPP, 1990, 22).
The time required for the completion of the bont
tick’s life cycle varied greatly, according to temperature and humidity. Lounsbury thought the minimum
period required was probably around 8 months, but
the process might take as long as two years if retarded by cool conditions. He was surprised by the
extraordinary vitality of the ticks, which were able
to survive on the pastures for six months without
feeding. The life cycle of the bont had obvious significance for techniques of attacking the tick on the
animal, for it was now understood that during a life
cycle which might take a year or 18 months to complete, the bont tick lived on its host for less than a
month (Lounsbury, 1899).
The link between heartwater and the bont tick had
not, however, yet been proven. Lounsbury commenced attempts to infect animals by means of bont
ticks in September 1899 and published important
results by mid-1900. In the Eastern Cape, the vets
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infected goats by inoculating them with blood taken
from an animal at the height of the disease. When
these goats became ill they were deliberately infested with bont ticks, which were then taken from Fort
Beaufort to Lounsbury’s Cape Town office, so that
the experiment, which was conducted “in an old
shed in the heart of Cape Town”, could be completed away from sources of accidental “natural” infection. When the nymphs had moulted into adults
they were placed on susceptible goats brought from
the heartwater-free district of Stellenbosch, several
of which soon became fatally ill (CPP, 1901, 16).
The vets used temperature charts and post mortem
evidence to confirm heartwater. As further proof,
the veterinary bacteriologist William Robertson
used their blood to reproduce the disease in other
goats. The red ticks, on the other hand, were apparently non-pathogenic, while a group of goats kept
clean of ticks remained perfectly healthy, a result
which corroborated the vets’ observation that the
disease was not directly contagious (AJCGH, 1900,
310). Lounsbury concluded that the bont tick was
“unquestionably” an agent in the transmission of
heartwater and that “intermittent parasites such as
ticks are the chief if not the sole agents is, it seems,
beyond question” (Lounsbury, 1900).
This was convincing evidence and Hutcheon, previously sceptical, now agreed that the bont tick was the
principal if not the only medium of communicating
heartwater to sheep and goats (CPP, 1901b). There
was a growing perception both among Eastern Cape
farmers and in the Department of Agriculture that
this was worthwhile and important research. This was
reflected in an increase in the scale of the heartwater
experiments in 1901, when the Cape government purchased 50 goats and funded an experimental station
at Rosebank, near Cape Town, with costs charged to
the veterinary vote (CPP, 1902, 44-5).
Hutcheon thought that it would be of great value to
know the relations between cattle, sheep, goats and
ticks in the propagation of the disease, for although
heartwater seemed to affect small stock exclusively,
the bont was primarily a cattle tick (CPP, 1901b, 8).
In giving evidence to the Stock Diseases Commission, the farmer John Webb had explicitly linked
heartwater to cattle. “Boschsickness” or “gallsickness” of cattle were also, he argued, caused by the
bont tick (CPP, 1877, 108-9). Lounsbury also noted
that Alexander Edington, the Cape’s government
bacteriologist, claimed to have produced a case of
heartwater in an ox by blood inoculation, which
suggested that the disease might occur naturally in
cattle (CPP, 1902, 35).
Hutcheon doubted that heartwater was a cattle disease, but nevertheless sent Lounsbury two calves for
transmission experiments. Lounsbury and the vets
began by infesting the calves with infected nymphs
brought from the Eastern Cape district of Somerset
East. Both of these animals became ill, but post-
mortem failed to reveal the characteristic lesions of
heartwater. The next phase of the experiment was to
determine if the disease could be transmitted from
the calves to susceptible goats. Larval bont ticks
(known to be uninfective) were fed on the sick
calves and blood was drawn at the same time. Both
the bite of nymph and blood inoculation proved
fatal to goats, but while the temperature curves
were characteristic of heartwater, the characteristic
post-mortem lesions were absent. The vets, however, found that they could produce heartwater in its
typical form by “passaging” the disease through several generations of goats by blood inoculation.
Lounsbury concluded that there was “no doubt
whatever that in every case the infection transmitted
was that of heartwater”, and that the disease could
be transmitted between calves and small stock by
the bont tick (AJCGH, 1902, 165-9;).
By the end of 1902, therefore, the role played by the
bont tick in the transmission of heartwater was substantially documented and proven. The collaboration of vets, entomologists and some progressive
farmers had confirmed a long-standing popular
belief about the nature of heartwater. But relatively
little was known about the aetiology of the disease
and germ practices (such as microscopic examination) failed to reveal a specific cause. From the public perspective, the most convincing feature of the
experiments was Lounsbury’s demonstration of the
transmission of heartwater by ticks. Thus the role
played by various environmental factors in the propagation and transmission of the disease was more
important than germ theory in suggesting methods
of prevention.
The experimental proof that ticks transmitted several stock diseases was a powerful influence on the
way vets, officials and farmers thought about prevention. Tick transmission explained much about the
apparent relation of certain diseases to climate, environment and locale. For farmers in the heartwater
area, Lounsbury’s work was authoritative. Ticks
were something that farmers could understand as a
factor in the incidence of disease. They were visible,
sometimes large and infested stock in enormous
numbers, while their blood-sucking habits and
vicious bite were an obvious source of harm to animals. From late 1902, officials in the Cape’s Department of Agriculture began, through the Agricultural
Journal, something of a “propaganda war” against
ticks, which were now accepted as the cause of the
collapse of the wool industry in the Eastern Cape.
Ticks had become “a terrible scourge” in parts of the
Colony, upon which they imposed a “peculiar form
of blood tax”. The significance of Lounsbury’s work
was described in terms of its potential economic
importance, for tick eradication was a means of:
Ultimately winning back the thousands of acres which
have been rendered useless for small stock owing to
the prevalence of this scourge …. That part of the
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country would be rehabilitated, and again become the
great meat and wool producer of South Africa
(AJCGH, 1902, 290-2).
By 1902 the time had come, according to the Agricultural Journal, when “the destruction of ticks
becomes as much an affair of national importance as
the eradication of scab” (AJCGH, 1902, 406). The
tick problem, it was argued, threatened both the
urban and rural communities because it limited pastoral production and reduced the food supply
(AJCGH, 1903, 235-6). The lead was taken from
both the USA and Australia, where, following Smith
and Kilborne’s discoveries, tick eradication was
being used as a method of preventing tick-borne diseases. In the USA, the construction of public and
private dipping tanks was subsidised from public
funds. Cape officials began to report these developments in the Agricultural Journal in 1896, a few
years before Lounsbury’s discoveries (Hutson, 1994,
93-5; Palladino, 1996).
The investigation of arsenic dips at the Cape was
particularly associated with the Adelaide dairy
farmer Llewellyn Roberts, whose farm, Cottesbrook, became a centre for dipping experiments in
the early 1900s. Roberts was convinced that if ticks
had ravaged stock farming in the eastern districts,
farmers could use technology to control them. In
1899 he read a paper to the Fort Beaufort and Adelaide Farmers’ Association. Adopting the moral tone
of a self-conscious “progressive”, he exhorted his
fellow farmers to take action:
Now gentlemen, you have got to face this. Are you
satisfied to sit still and gradually see your farms
ruined by these little pests? Will you be satisfied in
twenty years or less to give up your sheep and your
cattle for fruit growing as the Lower Albany farmer is
doing today (AJCGH, 1899, 371).
Roberts invited Lounsbury to Cottesbrook to devise
a means of destroying the large and resilient bont
tick. They experimented on a comparatively large
scale with 70 head of cattle, eventually working out
an arsenic dip at a concentration which would kill
the tick without apparent harm to the animal (CAD,
1899).
When “African coast fever”, later called East Coast
fever, broke out in the Transvaal in 1902, the control
of ticks came to the top of the pastoral agenda. This
proved to be a deadly tick-borne disease of cattle,
which threatened to destroy the pastoral industry
(Cranefield, 1992). Lounsbury’s demonstration that
the brown tick, a species common and widespread in
the Cape Colony, could transmit this disease did nothing to reassure farmers (CPP, 1904, 12-3). The Agricultural Journal noted that “uneasiness and anxiety”
were growing among farmers in the eastern districts
of the Cape as the spread of the disease in the Transvaal was reported. With a reported mortality rate of
above 90 per cent, the disease seemed frighteningly
virulent. It was feared that the African Coast fever
might work its way down from Natal and reach the
295
Eastern Province through the “native territories” of
the Transkei (AJCGH, 1903b, 385).
The “tick plague” was a major subject of discussion
at the annual Congress of the South African Agricultural Union in April 1903, which urged legislation for the compulsory eradication of ticks and
called on the Cape government to establish an
experimental farm to investigate methods of tick
eradication more fully (AJCGH, 1903b, 385). The
Cape government, however, was slow to act, and it
was not until 1904 that Hutcheon and Roberts
began experiments with a dipping tank at Cottesbrook (AJCGH, 1903a, 249-53). By then other
farmers were already conducting their own investigations. A Fort Beaufort farmer, Gordon Campbell
of Rocklands, completed a tank in May 1903. This
tank, which had cost about £ 100 to construct, was
claimed to be very effective at killing ticks and
could allegedly process 60 cattle in four minutes.
Others soon followed: in Bathurst, two farmers,
William Ford and Stephen Smith, began the construction of a cement and concrete structure at a
cost of £ 150; and at Kei Road, King William’s
Town, another farmer, Robert Warren was arranging
for the construction of a public dipping tank
(AJCGH, 1904, 7).
Although not all farmers were convinced of the benefits of arsenic dipping, a broad consensus was
emerging among farmers, officials and the government vets in favour of dipping against ticks. By
1909, the number of dipping tanks in the Colony
had increased to 175 (AJCGH, 1909, 472-3). Tanks
were also in use in Natal and Transvaal, where construction was greatly accelerated by the spread of
East Coast fever during the 1900s (Cranefield,
1992).
Heartwater as a germ disease
Following the British defeat of the South Africa
Republic (Transvaal) in the South African War
(1899-1902), Lord Milner began the economic and
political reconstruction of the Transvaal and the unification of the South African colonies. Milner was
committed to “constructive imperialism” and the
economic development of the region, including its
agriculture. He appointed Frank B. Smith to preside
over a substantial Department of Agriculture, a
large proportion of which was constituted by veterinary staff (Krikler, 1993, 78). Apart from field staff,
Smith recruited Arnold Theiler, a Swiss veterinarian
who had immigrated to the Transvaal in 1893, to
head a veterinary laboratory which was established
near Pretoria in 1902 (Gutsche, 1979; Theiler,
1971). Theiler transferred to modern facilities at
Onderstepoort, about ten miles north of Pretoria, in
1908. The laboratory formed the basis of the Onderstepoort Veterinary Institute, which controlled veterinary research for the whole of South Africa following the union of the southern African colonies in
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1910. The Onderstepoort Veterinary Research Institute achieved world renown during the 1920s and
1930s, when many important papers were published
on parasitology, bacteriology, plant toxicology,
immunology and nutrition studies (Brown, 2005).
As Director of Veterinary Services until his retirement in 1927, Theiler oversaw much important
research, which reflected the continuing importance
of pastoral production to the South African economy. In the context of a racially segregated society,
the research agenda was largely shaped by the concerns of European stock farmers. Up to the early
1910s, Theiler concentrated on the explication of
“tropical or sub-tropical diseases” of animals, which
he broadly defined as diseases that were transmitted
by arthropods (Transvaal Department of Agriculture
1909, 21). In this regard, research into East Coast
fever was particularly important and led to the
investigation of several other tick-transmitted diseases.
Theiler was interested in heartwater for two reasons. First, it was of economic importance in the
Transvaal. Merino sheep and angora goats could
not survive in the bushy lowveld because of their
vulnerability to the disease. Only the “common Kaffir goat” and fat-tailed sheep could be kept there,
probably because, Theiler thought, “they have
become immune over the long run of time”. More
importantly, he thought that heartwater was probably a lot more common in cattle and calves than
was generally acknowledged by farmers. In the
course of examining the corpses of experimentally
produced cases, he found that the lesion commonly associated with the disease – the filling of the
heart sac with liquid – was not very common in cattle. He believed that a lot of cases which farmers
attributed to the vaguely defined conditions “gallsickness”, “drunk gallsickness” and bushsickness,
were probably really heartwater (Theiler, 1904,
114-116 and 122).
Secondly, during the period immediately following
his appointment as Government Veterinary Bacteriologist, Theiler’s relation with researchers in the
Cape’s Veterinary Department, and more specifically with Alexander Edington, the Director of the
Cape’s Bacteriological Institute, had become competitive. Edington published an ill-advised article in
the Journal of Comparative Pathology and Therapeutics in which he claimed that African horsesickness, a deadly disease of equines, and heartwater
were the same disease occurring in different species
of animals (Edington, 1904). Theiler was quick to
use his own transmission experiments to demonstrate that horsesickness could not be produced in
sheep, goats or cattle. He published an article which
so convincingly contradicted Edington’s work that it
further undermined Edington’s already shaky reputation (Theiler and Stockman, 1905). The episode
contributed to the closure of Edington’s Institute,
and left Theiler as the uncontested leader in veterinary research in southern Africa.
Apart from this somewhat politically motivated publication, Theiler’s research on heartwater did not
add much to the knowledge previously gained by the
Cape government scientists. The government vet
R.W. Dixon demonstrated that heartwater could be
produced in susceptible animals by blood inoculation and passaged from animal to animal. Thus, by
the early twentieth century, heartwater had been
defined as a specific disease entity, although the
putative cause, a microbe, had never been identified
(Gilfoyle, 2003b). Theiler classified heartwater as
an inoculable disease caused by an “ultravisible
virus” and briefly tried to devise a prophylactic inoculation. The results obtained, however, were so
unpromising that he soon dropped this line of
research. The exigencies of the 1900s and 1910s,
notably the battle to contain East Coast fever,
pushed heartwater down the research agenda, while
the development of reasonably effective methods of
controlling the bont tick using arsenical dipping,
which had been developed in the Eastern Cape, rendered the problem less pressing.
Nevertheless, heartwater remained a disease of considerable economic importance on lowveld pastures
inhabited by A. hebraeum, which were otherwise
suitable for cattle farming. The disease took a heavy
toll of calves (heartwater was increasingly seen as
principally a disease of cattle), but it was also a serious problem for farmers who wanted to use imported breeding stock, which seemed to be highly susceptible to the disease. During the 1930s, officials
and farmers increasingly perceived heartwater as an
impediment to cattle farming in the Northern Transvaal bushveld. According to E.G. Hardy, the Union’s
Superintendent of Dairying, many ranchers and
dairymen were barely able to make a living there.
The problem, as Daly saw it, was that farmers were
unable to import bulls to upgrade stock, because
they were almost certain to die of heartwater.
Localised in-breeding over generations had caused a
deterioration of the quality of stock and low productivity in terms of meat and milk (AOVI, 1934).
During the mid-1930s, when East Coast fever had
been confined to a relatively few infected areas, the
veterinary bacteriologist E.M. Robinson, noted that
“heartwater is perhaps the most serious disease of
cattle and sheep in the Union” (AOVI, 1935). Writing in the 1940s, Petrus J. du Toit, who replaced
Theiler as Onderstepoort’s Director in 1928, identified heartwater as
“Today the most serious problem cattle farmers have
to contend with in the low-veld areas of South
Africa….. It forms a serious and almost insuperable
obstacle against cattle improvement and successful
cattle farming generally in the areas where it is
enzootic” (du Toit, 1945).
A major revival of heartwater research during the
mid-1920s, however, depended on American inter-
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ests. Scientists at the Rockefeller Institute in the
United States had been researching a dangerous disease of humans and animals called Rocky Mountain
spotted fever (Harden, 1990). Theiler had apparently read about this disease, which was also transmitted by ticks and was symptomatically similar to
heartwater. He therefore contacted Simon Flexner,
the Director of the Rockefeller Institute, and invited him to send a scientist to study heartwater in
South Africa. Flexner seems to have agreed that
heartwater studies might throw light on the nature
of Rocky Mountain spotted fever, for in 1924, he
seconded E.V. Cowdry to Onderstepoort (AOVI,
1924). By the mid-1920s, a small group of diseases
(Rocky Mountain spotted, typhus and trench fever)
were attributed to rickettsias, small organisms
which were like bacteria in appearance, but which,
unlike bacteria, inhabited cells. Rickettsial diseases
were thought to be non-contagious, but generally
transmitted through the bite of arthropod species.
Cowdry had already completed a study of Rocky
Mountain spotted fever and was experienced in the
laborious task of searching for the rickettsias in tissue sections.
Once Cowdry had succeeded in establishing a reliable source of material by passaging the infection
through live animals, he achieved an immediate success. Nothing was to be found by examining fresh
blood smears, but the persistent searching in sections
of the kidney (most commonly), spleen, lymph
glands and parts of the brain revealed small, coccuslike bodies in the cells lining the capillaries (endothelial cells) of these organs. They were similar in size,
shape and staining characteristics to the rickettsias of
typhus and Rocky Mountain spotted fever. Within
the endothelial cells, the organisms were grouped in
clumps, apparently multiplying by simple division.
Cowdry took considerable pains, particularly in the
use of staining techniques, to prove that these were
really foreign bodies rather than cellular components
or the products of cell degeneration. There was no
evidence, however, that these organisms produced
any specific damage to the cell, except that in some
cases their multiplication caused the cells to become
distended and even to burst, releasing the organism
into the circulation. There was nothing to indicate
that the organisms spread to neighbouring cells. The
presence of the organisms was closely associated
with the course of fever in the infected animal. They
appeared shortly after the onset of fever and began
to decline in numbers after the peak of the fever, persisting in the tissues for a few days following the disappearance of clinical symptoms. In spite of the correlation of the appearance of organisms with the
clinical symptoms, there seemed no obvious way in
which they were connected with, or caused these
symptoms, or indeed the characteristic post-mortem
lesions (Cowdry, 1926a).
Cowdry complemented his work on heartwater in
animals with a study of the organism in ticks, to
297
demonstrate that it could always be found in ticks
which were able to transmit the disease, thus completing his “aetiological proof”. The picture in
nymphs and adults of A. hebraeum that had fed on
a sick animal in a previous stage of the life cycle was
rather complicated by the presence of several kinds
of micro-organisms. Nevertheless, he saw morphologically similar organisms in cells lining the intestine of the tick. As he had found in the sheep, the
organisms clumped together, sometimes distending
the cell to the point of rupture and thereby escaping
into the contents of the intestine. Unlike in some of
the protozoan diseases, in which the parasite was
found in the salivary glands of the tick, Cowdry
thought that the causal organism of heartwater was
probably transmitted by regurgitation. The infectivity of the ticks seemed to depend on the presence of
these organisms, for the symptoms of the disease
only appeared when ticks containing the organism
fed on susceptible animals. Generally, ticks in which
the organism could not be detected failed to transmit the disease (Cowdry, 1926b).
Cowdry argued that this proved the organism was
the cause of heartwater because:
(1) It could be detected in cases of heartwater and
its appearance and disappearance was closely
associated with the development of fever and
the infectivity of the blood. They were absent in
healthy animals
(2) Similar organisms could be found in ticks which
had fed on sick animals, but they could not be
detected in controls which had fed on healthy
animals.
(3) The symptoms of heartwater appeared when
ticks with the organisms in their alimentary tract
fed on healthy animals, but never when in animals which were infested with ticks in which the
organisms were not present.
(4) The “cycle” could be completed by demonstrating the presence of the organism in animals
which had become sick following the bite of an
infected tick (Cowdry, 1926b, 181-98).
The requirements for “proof” had relaxed since the
more rigorous requirements of Koch’s postulates,
which insisted on the cultivation of the causal
organism outside the body. Proof here consisted of
little more than the observation of a consistent and
close association between the presence of the organism and the symptoms of the disease. Furthermore,
Cowdry did not speculate in the published material
about the processes by which the organism acted
upon the body to produce the very striking and
severe symptoms of heartwater. He was more forthcoming, however, in classifying the newly discovered
organism. As mentioned above, heartwater seemed
to have much in common with Rocky Mountain
spotted and typhus, which were attributed to
species belonging to the genus rickettsia. In his own
definition of the rickettsias, Cowdry stressed “the
ability of the organisms to lead an intercellular (sic)
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existence, their location in the tissues, their host
specificity, their gram-negative properties, and their
bacterium-like morphology” (Cowdry, 1926a, 171).
To this definition Cowdry added another set of specifications by M. Hertig and S.B. Wolbach, contemporary experts on typhus fever, that rickettsia were
“Gram-negative, intracellular, bacterium-like organisms found in arthropods” (Hertig and Wolbach,
1924). As the organism of heartwater fitted all of
these requirements, Cowdry suggested that it should
be classified as Rickettsia ruminantium.
Although Cowdry stayed at Onderstepoort for less
than a year, collaboration with the Rockefeller Institute proved fruitful during the late 1920s. Theiler
and Du Toit provided further evidence supporting
Cowdry’s conclusions on heartwater by demonstrating that heartwater could be transmitted by the
injection of ground up nymphs of A. hebraeum
which had recently fed on an infected animal.
Cowdry published a series of articles on heartwater
and East Coast fever in the American Journal of
Experimental Medicine. These articles publicised
the problems of stock diseases in southern Africa
(and Africa more generally) while acknowledging
work already done at Onderstepoort. The Rockefeller collaboration was part of the continuing integration of Onderstepoort into international research
networks in a variety of areas such as protozoology,
nutrition and toxicology.
It was, however, scientific research which had little
immediate practical value for prevention. In this
regard, there was little advance on Spreull’s 1904
definition of heartwater as
A specific febrile disease affecting sheep, goats and
cattle, in South Africa and due to an ultra-visible virus
transmitted by the bont tick, Amblyomna hebraeum
(Koch) (Spreull, 1904, 433-442).
In a major review of the heartwater problem published in 1931, the Onderstepoort scientist Raymond Alexander advised that “with our present
knowledge the only rational prophylactic of any
importance is the elimination of infected ticks”
(Alexander, 1931). This advice was based upon
knowledge about the bont tick gleaned by Charles
Lounsbury in the 1900s.
Nevertheless, Cowdry’s research stimulated a series
of studies on the aetiology of heartwater from the
late 1920s. One of the major problems presented by
the disease was the connection between the apparently innocuous presence of R. ruminantium and
the dramatic symptoms. Further histopathological
studies could only confirm the presence of the rickettsias in the lining of blood vessels in kidney, cerebral cortex and various other organs and reveal a
reduction in the number of macrophages (one of the
categories of white blood cell) (Steck, 1928). Any
direct relation between the apparent aetiological
agent and the pathology of the disease remained
obscure. It seemed that some factor other than the
rickettsias must be at play. Raymond Alexander,
considering all the evidence available in the early
1930s, concluded that:
It seems most likely that the alterations are due to a
noxe which is spread diffusely by he blood stream. It
must remain for future investigation to determine the
nature of that noxe (Alexander, 1931, 109).
This distinction between R. ruminantium in the
endothelial cells of the blood vessels and a “toxin”
or “virus” in the blood, which caused the physical
damage, raised questions about Cowdry’s perhaps
too easy aetiological conclusions, which were based
solely on the consistent presence of the organisms in
heartwater cases and infective ticks. The nature of
this “noxe” was the subject of further studies by
another distinguished Onderstepoort scientist
Willem Neitz, assisted by the technological innovations of a colleague, Charles Jackson. Jackson
devised a means of taking “intima smears” from the
endothelial lining of the jugular veins of infected
sheep, a tissue in which the rickettsia were found to
be particularly numerous (Jackson, 1931).
Using Jackson’s techniques, Neitz was able to isolate
the rickettsias by dissecting out the jugular vein of
an animal in the throes of the disease and washing
them out to remove all traces of blood. The veins
were pinned out flat to expose the inner surface and
finely scraped out to retrieve the endothelial cells
which contained the organism. Injections of emulsions of these cells produced the typical symptoms
of heartwater with the incubation period in a high
proportion of susceptible sheep, convincing evidence that the rickettsias were in some sense the
direct cause of the disease. Neitz also cast doubt on
the theory that death in heartwater was caused by a
toxin circulating in the blood. Massive blood transfusions from sick into healthy animals did not
appreciably hasten the course of the disease, an
unlikely result if a toxin was the cause of the symptoms (Jackson and Neitz, 1931, 49-52).
Neitz’s microscopic examination of various tissues
was revealed an extremely confusing picture. Rickettsias in the endothelial cells revealed a variety of
sizes and shapes, included ring, horseshoe and
“pleomorphic” entities. This variety suggested the
possibility of a life cycle similar to that of protozoa,
possibly with an ultravisible phase circulating in the
blood which was instrumental in producing the
symptoms and lesions. Against this had to be
weighed earlier evidence amassed by Alexander that
the causal agent of heartwater did not pass bacteriological filters and seemed to be closely associated
with the blood cells, from which it could not be
detached by washing (Alexander, 1931, 97). Neitz
also identified rare single structures which he took
to be rickettsias circulating freely in the blood. Even
then, however, he had to keep in mind the possibility that these might be “artefacts” released during
the process of taking blood (Jackson and Neitz,
1931). Neitz suspected that the freely circulating
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forms played an important role in aetiology, but was
unable to come to a definite conclusion. He was also
unable to make use of important technical innovations in the propagation of viruses and rickettsias in
small laboratory animals and, in the chicken
embryo, which enabled rapid advances in the study
of many diseases during the 1930s and 1940s
(Mason and Alexander, 1940). Unfortunately for
Neitz and his colleagues, heartwater proved refractory to these methods.
The problems of aetiology in heartwater proved
insuperable during the 1940s. In a general survey of
1949, the veterinary scientist M.W. Henning was
able to do no more than speculate that the organism
was probably attached to the red blood cells in such
a way that it could not easily be removed or seen
(Henning, 1949, 826). Instead, research into heartwater took a more applied turn, as scientists at
Onderstepoort concentrated on devising a means of
immunising animals against the disease. Nevertheless, the study of the biology of R. ruminantium and
aetiology of heartwater was an example of the extension of the tropical diseases model, with its emphasis on arthropod vectors, environment and climatic
factors, into the field of rickettsial diseases.
The investigation of immunisation
During the rinderpest epizootic of 1896-98, veterinary scientists in southern Africa developed methods of immunising cattle by injecting them with
infective blood and the blood serum of recovered
animals (Gilfoyle, 2003a). As these veterinary scientists believed that these methods achieved a
degree of success, they sought to extend them to
other diseases. Theiler tried injecting recovered
sheep with blood taken from animals with symptoms of heartwater, hoping that injections with
blood from different species (sheep, goats and cattle) might produce a particularly effective serum.
This “hyperimmunised” serum seemed to protect
against injections of infective blood but failed to
produce a longer term immunity. Experiments of a
similar kind by a government vet, James Spreull, in
the Eastern Cape produced equally negative results
and were abandoned by 1905 (Theiler, 1905).
A major practical problem for Theiler was that
blood taken sick animals soon lost its infectivity.
The causal agent of heartwater or “virus”, as Theiler called it, was extremely fragile in the blood once
it had been removed from the sick animal. This presented considerable difficulties for experimental
research, because infective material could not be
stored for any length of time. Theiler thought that a
vaccine was impossible given the characteristics of
the “virus” and “had little hope that the difficulty
[could] ever be overcome” (AOVI, 1921). Veterinary advice on heartwater prophylaxis during the
1910s and 1920s thus continued to emphasise the
value of arsenical dipping, although Theiler admit-
299
ted that the bont tick, which was resilient and
mobile, was one of the most difficult species to
eradicate. As recovered animals seemed rapidly to
lose their capacity to infect ticks, the dipping and
removal of animals from an infected area was also
an effective means of stopping an outbreak (Theiler
1909; Spreull 1922).
The possibility of an effective prophylactic was not
seriously raised again until the mid-1920s, during
Cowdry’s visit to Onderstepoort. In 1925, Petrus du
Toit and Raymond Alexander instigated a series of
experiments to attenuate the “virus” of heartwater
by passaging it through a lengthy series of sheep. He
hoped that the “virus” might become sufficiently
attenuated to produce a vaccine, as had been the
case in some other diseases. Passaging entailed a
laborious process of subinoculation of blood from
sick to healthy animals in a continual series, under
controlled conditions to avoid concurrent infection
from ticks. After many passages over a period of
five years the organism was still fully virulent and a
vaccine as elusive as ever (du Toit and Alexander,
1931, 151). At this time, Max Theiler, Arnold’s son,
who was working in the United States, succeeded in
attenuating yellow fever virus by passaging through
the brains of mice. Alexander was excited by this
discovery and made similar experiments with heartwater, but again with no success (Alexander, 1931).
During the mid-1930s, Wilhelm Neitz set out to
question some of the assumptions previously made
about immunity in heartwater, for example, the possibility of immunogenically different strains, which
would present serious problems for any future
means of immunisation. Neitz collected ten “strains”
of heartwater from farms in different parts of the
Transvaal for use in a long-running series of crossimmunity tests. He found that although the strains
varied considerably in virulence, sheep which had
recovered from one strain showed a strong immunity to all the others. This was a significant result,
because it meant the any vaccine would need to
incorporate only one strain, unlike other diseases
such as horsesickness, for which effective vaccines
needed to be polyvalent (Gilfoyle, 2006). Neitz also
found that immunity in recovered sheep was sometimes partial and they could occasionally be reinfected without showing clinical symptoms. This was
significant for control because apparently immune
sheep might act as “reservoirs” capable of infecting
ticks (Neitz, 1939).
Neitz made some important practical discoveries
during the late 1930s. He found that very young
calves, up to around three weeks old, were highly
resistant to heartwater and could be injected relatively safely with infective sheep’s blood. He also
found he could treat heartwater induced by injections of infected blood with the sulpha drug, Uleron,
which was marketed by the German Bayer Pharma
company. In the course of the Second World War,
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Uleron went off the market, but Neitz later found
that he could use other drugs of similar chemical
composition, such as Solupyridine manufactured by
May Baker and Sulphamethazine from African
Explosives (AOVI, 1947). Neitz’s laboratory results
were confirmed by some small-scale field experiments carried out around Potgietersrus in the Northern Transvaal during 1940. Significantly, from the
point of view of prophylaxis, he found that animals
which recovered from the infection through the
administration of drugs were apparently immune to
further infection (Neitz and Alexander, 1941). Given
that different strains of heartwater were apparently
immunogenically similar, these findings opened the
possibility of producing immunity by using sulpha
drugs to control an induced infection.
During the 1930s, officials perceived heartwater to
be a particularly serious problem in the bushveld of
the Northern Transvaal. E.G. Hardy, the Union’s
Superintendent of Dairying, reported that cattle
farmers and dairymen were barely making a living
here. The problem, as Hardy saw it, was that farmers
were unable to import bulls to upgrade stock,
because they were almost certain to die of heartwater. Local cross-breeding over generations, he argued,
had caused a degeneration of the quality of stock and
low productivity in terms of meat and milk.
The problems of Northern Transvaal stock farmers
lay behind the establishment of the government
farm “Mara” in the mid-1930s for the breeding of
“acclimatised” bulls. But Neitz and his colleagues
were able to use these concerns of farmers to negotiate a more extensive trial of the “infect and cure”
method of immunisation. J.M. Marks, the lands
manager of the African and European Investment
Company, which had large estates around Warmbaths in the Northern Transvaal, was interested in
trying out Neitz’s method. The management of the
estates, which supplied mining compounds with
beef, had found that in spite of regular five-day dipping, losses from tick-borne diseases were still
enough to prevent profitable beef production. Furthermore, the continual interruption of grazing
entailed by dipping caused a loss of weight which
counterbalanced the potential benefits. The Company offered Neitz and Alexander the opportunity of
an extensive trial of his method, which ran for three
years from 1939 to 1942 (Neitz, 1945, 138).
The experiment was an interesting one for the scientists, not least because it gave them insight into
the mortality caused by tick-borne diseases under
ranching conditions. Although susceptible cattle
introduced to the estate contracted protozoal diseases, particularly redwater fever, these diseases
rarely caused death in cattle born and bred on the
estate. Heartwater, however, was much more common and frequently deadly. It appeared that a state
of “endemic stability” could be achieved against redwater fever, but that heartwater remained a serious
problem and was the chief cause of mortality under
ranching conditions. In 1939, Neitz and Alexander
began a controlled experiment entailing the immunisation of grade (cross-bred) Aberdeen Angus
calves. Mortality in the control group of 195 nonimmunised calves was so high that, in the third year
of the experiment, the management of the estate
became alarmed and decided to sell most of the
herd on, as it looked as though most of these animals would eventually die from heartwater. In contrast, mortality in the immunised herd of 1,321 animals was not more than five per cent. The problem
was that the process of immunisation had killed a
similar number, raising the overall mortality rate to
close to ten per cent (Neitz, 1945, 143-47).
The experiments were considered a qualified success. Marks was sufficiently impressed to have the
estate manager continue with immunisation, which
required the maintenance on the estate of the heartwater strain in susceptible sheep by passaging. Du
Toit was cautious about publicising details of these
experiments after they were wound up in 1943. He
feared the method was too dangerous for general
use and wished to discourage requests for the mass
immunisation of young cattle (AOVI, 1942).
The scientists believed the control of heartwater in
domestic animals depended upon “the permanent
rupture of at least one link in the chain of cyclical
development of the disease”. The possibilities were
the elimination of the tick vector, the elimination of
disease “reservoirs” or the replacement of susceptible
hosts with immune or resistant animals (Neitz and
Alexander, 1941). They saw arsenical dipping as a
means of limiting tick infestation to reasonable levels, rather than of complete eradication. Nor was this
necessarily desirable, as some degree of contact with
ticks was necessary to maintain resistance against
protozoal diseases, such as redwater fever. The elimination of the disease reservoir was also problematic, given the discovery of R. ruminantium in wild
animals, while further research confirmed domestic
animals as reservoirs of infection (Neitz, 1944).
Neitz and his colleagues aimed at exploring in more
detail the duration of immunity in recovered sheep.
The technique was to inject recovered (“immune”)
sheep with infected blood, and then to test the presence of “virus” circulating in their blood by a series
of subinoculations over specified intervals into susceptible sheep. To their surprise, they found that the
blood of recovered (in effect, immunised) merinos
was sometimes infective for susceptible animals for
lengthy periods. The agent of infection was apparently circulating in the blood of the “immune” animals, even though no clinical reaction was detectable
(Neitz, Alexander and Alelaar, 1947).
These were striking results, suggesting that immunity against heartwater was in some ways akin to “premunity” in the protozoan diseases, rather than
immunity in the bacterial diseases, such as anthrax,
26-08-2009
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Pagina 301
in which the bacilli were rapidly destroyed. These
experiments supported previous observations that
heartwater sometimes affected sheep and goats on
farms where the disease had apparently been absent
for a long time. From a practical point, however, it
meant that sheep could themselves provide a possibly permanent reservoir of infection which could
not be eliminated. Vets thus came to see immunisation as the most promising means of preventing the
disease in susceptible animals (Neitz, Alexander and
Alelaar, 1947).
The major practical problem for the extension of
immunisation against heartwater, apart from the
danger inherent in the process itself, was the very
limited viability of the “virus” once blood was
removed from the infected animal. The blood had to
be used within 24 hours of drawing. Heartwater
vaccination, however, soon began to attract some
public attention. Farmers wanted Onderstepoort to
supply them with sheep infected with a mild strain
so that they could immunise their own bulls without
using drugs. As Onderstepoort’s Director, du Toit
was unwilling to agree to this, foreseeing that it
would entail the despatch of thousands of infected
sheep every year. The practical problems inherent in
this method were revealed in 1944, when Onderstepoort supplied a sheep infected with a mild strain
to the Swaziland Veterinary Officer, W. Barnard.
The intention was that Barnard should maintain the
strain (designated Ball III) by subinoculating a series
of susceptible merinos so that farmers could bring
in their bulls whenever immunisation was required.
In practice, however, Barnard found the matter surprisingly difficult. The “reservoir” sheep had to be
kept absolutely clear of bont ticks to prevent the
contamination of the pure strain, which Barnard
soon lost. He suggested that Onderstepoort send
him the required sheep, which could be ordered
well in advance, but du Toit was determined not to
set a precedent (AOVI, 1944).
During 1945, the vets began treating calves in the
immediate vicinity of Onderstepoort and soon afterwards the scheme was extended to farms within a
200-mile radius of the laboratory. Owners were simply expected to turn up at Onderstepoort on the
morning that the “reservoir” sheep were to be bled.
They were allowed to bring in calves for veterinary
treatment or to take the blood away to do the injections themselves. Later, farmers were invited to drive to Onderstepoort with a sheep for infection with
the mild strain, which they could later use to immunise calves (AOVI, 1949). Apart from the innate
difficulty of this method of distribution, it was also
extremely limited in its reach. Cattle owners in other parts of the country affected by heartwater, the
farther reaches of the Northern Transvaal and parts
of Natal and the Eastern Cape, which were far from
Onderstepoort, began to demand access to immunisation (AOVI, 1950).
301
The management of Onderstepoort initially refused
to consider supervising heartwater centres in other
parts of the country, hoping the extension of heartwater immunisation could be achieved by local farmers’ associations or similar organisations (AOVI,
1945). Initially, Onderstepoort scientists worked
with “progressive” farmers, such as J.S. Sprigg from
Alexandria (a breeder of Jersey cattle) and E.A.
Galpin from Naboomspruit, with whom they had
good working relations. The process of maintaining
the strain in practice was rather onerous. Because
infective blood needed to be used within 12 hours of
withdrawal, Alexander despatched it to the Eastern
Cape by airplane. Sprigg, for example, was required
to turn up promptly at Port Elizabeth airport armed
with a syringe and a sheep for immediate injection.
He then had to subinoculate another sheep approximately every 13 days in order to maintain the strain.
Sprigg also had to ensure that the sheep used were
not carrying a latent infection by keeping them completely free of bont ticks, itself no easy matter in a
low-lying district like Alexandria. Farmers issued
with blood evidently found it difficult to maintain
the strain (AOVI, 1951).
In the early 1950s, Alexander found that the “shelflife” of the blood could be substantially lengthened
by freezing and storing in dry ice. This obviated the
difficulties of supplying blood to farmers in the
Transvaal. The Eastern Cape, however, lacked the
facilities for producing dry ice. In 1952, Onderstepoort eventually bowed to the pressure exerted
by Eastern Cape farmers and agreed to the establishment of “heartwater stations” at East London
and Grahamstown under the supervision of government veterinary staff. In spite of persistent requests
for another station at Port Elizabeth, actual demand
for heartwater blood remained relatively small, and
had not exceeded 10,000 doses by 1954. The failure
of farmers to make full use of the new facilities
annoyed Alexander, who calculated that the vaccine,
which sold at one shilling per dose, cost close to six
shilling to produce. It was, he argued, a substantial
government subsidy to farmers.
Given the considerable difficulty in obtaining the
product, the necessity for close supervision and the
possibility that injection with drugs might be necessary, it seemed unlikely that immunisation against
heartwater would spread far beyond wealthier stock
farmers dealing with comparatively expensive animals. In an age in which the safety and efficacy of
vaccines could be measured with some accuracy and
some disease-producing micro-organisms could be
reliably attenuated according to a variety of methods,
heartwater immunisation was in many ways unsatisfactory. As Alexander put it, the injection of infected
blood was “scientifically a poor method”, not least
because if was difficult to guarantee the efficacy of a
product which was apparently in a state of deterioration from the time it was taken from the donor animal (AOVI, 1950). Heartwater presented extremely
302
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difficult problems to veterinary researchers during
the 1940s. It outlined the limits of scientific knowledge and techniques at a time when considerable
advances were being made in other fields and against
other diseases. For most farmers, the use of a variety
of environmental controls, especially dipping, which
had been in use since the late 1890s, remained the
most important means of controlling this damaging
disease.
Conclusion
Heartwater, a fatal disease of cattle and small stock,
had serious economic consequences for farmers in
parts of southern Africa. During the late nineteenth
century, British-trained veterinary scientists employed
by the Cape government were unable to offers farmers an explanation of disease. The American entomologist, C.P. Lounsbury, in using scientific method
to work out the transmission of heartwater, confirmed the beliefs of many farmers. Farmers were also
instrumental in adapting methods of control, particularly the dipping of animals in arsenical preparations,
which had been developed in American and Australia. During the 1920s, the American microbiologist
E.V. Cowdry used insights gained from his study of
Rocky Mountain spotted fever in the United States to
discover the micro-organism which caused heartwater. During the 1940s, South African veterinary scientists, who had gained qualifications at Onderstepoort, investigated techniques of proplylactic inoculation against heartwater. They achieved some success, but there remained considerable practical difficulties with these techniques. Farmers played an
important role in field trials of prophylaxis and
obtaining knowledge about the incidence of the disease under natural conditions. Methods of killing
ticks developed in the United States during the late
nineteenth century and adapted by local farmers
remained, however, the chief means of controlling
heartwater. The investigation of heartwater entailed
interactions with scientists in other parts of the
world, but the linkages were with the United States
and Australia, rather than with the imperial centre.
Acknowledgements
I would like to than the Economic and Social Research Council and the Wellcome Trust for grants which enabled me to
complete the research for this article.
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Veterinary Entomology, colonial science and the challenge
of tick-borne diseases in South Africa during the late nineteenth
and early twentieth centuries1
K. Brown
Wellcome Unit for the History of Medicine, University of Oxford, Oxford, UK.
Abstract. This article provides an historical overview of developments in veterinary entomology during the
late nineteenth and early twentieth centuries. During that period state employed entomologists and veterinary scientists discovered that ticks were responsible for transmitting a number of livestock diseases
in South Africa. Diseases such as heartwater, redwater and gallsickness were endemic to the country.
They had a detrimental effect on pastoral output, which was a mainstay of the national economy. Then in
1902 the decimating cattle disease East Coast fever arrived making the search for cures or preventatives
all the more urgent. Vaccine technologies against tick-borne diseases remained elusive overall and on the
basis of scientific knowledge, the South African state recommended regularly dipping animals in chemical solutions to destroy the ticks. Dipping along with quarantines and culls resulted in the eradication of
East Coast fever from South Africa in the early 1950s. However, from the 1930s some ticks evolved a resistance to the chemical dips meaning that diseases like redwater were unlikely to be eliminated by that
means. Scientists toiled to improve upon existing dipping technologies and also carried out ecological
surveys to enhance their ability to predict outbreaks. Over the longer term dipping was not a panacea
and ticks continue to present a major challenge to pastoral farming.
Key words: ticks, South Africa, East Coast fever, redwater, heartwater, dipping, ecological surveys, veterinary science, entomology.
Historiographical context
Over the last ten years or so there has been a
marked increase in the historiography of veterinary
medicine in South Africa. Earlier references to veterinary interventions, dating from the 1970s,
focused on African resistance to state attempts to
“stamp out” diseases such as the highly contagious
cattle diseases, rinderpest and tick-borne East Coast
fever (van Onselen, 1972; Bundy, 1987; Phoofolo,
1993). At the height of apartheid, historians wished
to show that African resistance to all forms of colonialism, including veterinary medicine, had a history that predated 1948, when the Nationalist Party
came to power. It is only in the aftermath of
apartheid, in the late 1990s, that historians have
begun to explore veterinary science on its own
terms. Since then a number of historians have
looked at the origins and development of the Cape
Veterinary Department from the 1870s, as well as
the expansion in bacteriological research at Grahamstown in the Cape and at the Onderstepoort
Veterinary Institute, near Pretoria, which became
the main centre for veterinary research in South
Africa, after it’s founding in 1908 (Beinart, 1997;
Correspondence: Karen Brown, Senior Research Officer, Wellcome Unit for the History of Medicine, University of Oxford,
45-47 Banbury Road, Oxford OX2 6PE, UK, Tel +44 (0)
1865 274616, Fax +44 (0) 1865 274605. e-mail: karen.
brown@wuhmo.ox.ac.uk
1 The quotation comes from Lounsbury’s speech to the
British Association for the Advancement of Science, 1905.
Gilfoyle, 2002; Madida, 2003; Brown 2005). In
addition, there have been studies of a number of
vaccines, developed in South Africa, that contributed to the decline in livestock mortality from
contagious and infectious diseases (Gilfoyle, 2003a,
2006a, 2006b).
Veterinary entomology has appeared in this literature in a number of contexts. In particular, in
accounts of vaccine research against arthropodborne diseases such as horsesickness, as well as
studies of the ecological inter-relationship between
arthropods and the wider environment that enabled
infections, such as nagana (or bovine trypanosomiasis, now trypanosomosis) to thrive (Gilfoyle, 2006a;
Brown, 2008a, 2008b). In relation to ticks-borne
diseases, work has covered early investigations into
heartwater (Cowdriosis), a disease that affects
sheep, goats and cattle (Gilfoyle, 2003b), as well as
efforts to eradicate East Coast fever (Theilerosis) in
bovines. Paul Cranefield’s book, entitled Science
and Empire: East Coast fever in Rhodesia and the
Transvaal, is the most comprehensive work on a
tick-borne disease (Cranefield, 1991). Cranefield
discussed the nature of colonial science as played
out in South Africa, together with the role of the
state in combating this infection. Controlling tickborne diseases helped to define the function of the
South African state, as governments assumed the
right to legislate on the management of private
farms in the interests of the national good.
This paper expands upon the existing insights into
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K. Brown - Ticks and Veterinary Entomology in South Africa
veterinary entomology in late nineteenth and early
twentieth century South Africa, by examining a
range of problems surrounding disease control, as
well as environmental explications of livestock
infections. The aim is to give a general overview of
veterinary entomology in relation to ticks rather
than to explore a particular disease. This contrasts
with the approach of Gilfoyle and Cranefield who
have looked specifically at heartwater and East
Coast fever respectively. These writers concentrated
on the Cape and the Transvaal and tended to overlook Natal, which also contributed to the growing
body of knowledge about these types of infection.
Farmers and scientists in Natal were notably active
in developing methods of reducing tick numbers by
dipping livestock in chemical solutions. After 1900
dipping became the major tactic for tick control. In
addition, neither historian focused on later research
into the ecology of ticks or the problems that
evolved in relation to dipping. As this paper hopes
to show, discovering the vector of a disease was a
difficult and time-consuming process, and even if
scientists found ways of mitigating the spread of
tick-borne infections, solutions could be relatively
short-term. By the 1940s scientists began to reappraise their existing understandings of tick-borne
diseases in the absence of effective and safe vaccines, compounded by the recognition that dipping
alone might not be the long-term answer because
some ticks had developed a resistance to the chemicals. Around the same time, ticks featured in South
Africa’s Zoological Survey, which examined how
animals interacted with the environment, partly
with the aim of improving scientific understanding
of the epidemiology of a number of diseases in
which wildlife and arthropods served as the reservoir or vector (Kolbe, 1982; Bigalke and Skinner,
2002). This demonstrated the growing complexity
of veterinary entomology: not only did research
include microbiological and chemical work, but also
forays into the environmental sciences. Veterinary
science in South Africa during the inter-war years
developed a strong environmental sense, revealed by
concurrent studies into the biology and ecology of
tsetse flies that transmit trypanosomosis, as well as
investigations into the propagation of toxic flora and
nutritional deficiencies in the veld (Gilfoyle, 2003c;
Brown 2007, 2008b).
The rural economy and the start of
Veterinary Entomology in South Africa
Veterinary entomology had established itself as an
important aspect of veterinary research by around
1900 because so many livestock infections were
transmitted by a range of arthropods, including ticks
and tsetse flies. Farmers and scientists also assumed
that diseases such as horsesickness and bluetongue
(a sheep disease) were conveyed by biting flies,
although the exact identity of the vectors remained
elusive until the 1990s (Brown, 2008a). Some
arthropods did not convey disease, but were parasitic, such as the acari mite, which caused a condition known as scab by boring into the flesh of sheep
and goats, thereby destroying the fleece (Tamarkin,
1999). The destructive impact of arthropods was
significant as livestock were important to the South
African economy, not only as a source of food, but
also fibres. During the nineteenth century, the Cape
emerged as one of the largest producers of wool in
the world, competing with Australia and New
Zealand for dominance of the global markets. In
fact, income from wool often outstripped that from
diamonds (discovered in the Cape in 1866) and
from gold, which did not really take off as South
Africa’s major export until the twentieth century.
Farmers imported merino sheep form Europe in
order to enhance their yields. These introduced
sheep were particularly susceptible to tick-borne
diseases in Africa as they had had no prior exposure
to them. The same was true of goats. Farmers also
introduced angora goats from the Middle East, and
by 1914 South Africa had replaced Turkey as the
primary producer of angora wool. Cattle were bred
less as an export commodity and more as a source
of meat and milk for the mining compounds and
cities that expanded exponentially during the twentieth century. Concerns about tick-borne and other
diseases precipitated the establishment of the first
veterinary departments in South Africa, in Natal in
1874 and the Cape in 1876 (Beinart, 1997, 2003;
Gilfoyle, 2002).
Racism in all the South African states meant that
the primary purpose of these veterinary departments
was to improve the economic standing of white
farmers. By the mid-nineteenth century, European
settlers had appropriated much of the land in South
Africa, increasingly forcing blacks to reside in what
became the African reserves. Some blacks were
allowed to live on settler farms as tenants, or else
they were employed as labourers. As the nineteenth
century wore on, it became increasingly difficult for
Africans to produce foodstuffs and fibres for a commercial market as they lacked access to land and
capital. Many blacks were subsistence farmers for
whom livestock, especially cattle, had a cultural
rather than an economic value. African societies
were highly patriarchal, and men exchanged bovines
as bridewealth for wives. A man who owned a large
number of cattle had power and status within his
community. Disease undermined the ability of
Africans to increase their flocks and herds also,
although their economic ethos could be different to
that of white settlers (Bundy, 1979).
In the 1870s a white, self-styled “progressive”, farming elite who wanted to reduce livestock mortality
and improve their capacity to compete on the international markets, clamoured for the establishment
of scientific veterinary departments in the Cape and
Natal. This came in the wake of changing understandings and developments in western medicine.
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Work in France and Germany, in particular, by
Louis Pasteur, Robert Koch and their acolytes led to
the gradual dominance of “germ” theories and the
possibilities that diseases could be controlled if scientists could isolate the specific microbe and develop a vaccine to prevent susceptibility to a given
infection (Brock, 1988; Geison, 1995; Worboys,
2000). Twenty years later, demands for state entomology departments also came in the wake of discoveries elsewhere. In the Cape, the most powerful
lobbyists were wine producers, who saw their vineyards rot in the wake of phylloxera (Lounsbury,
1940; Brown 2003a). Nevertheless, ticks were also
a problem and the Cape’s first official entomologist,
the American Charles Lounsbury, was especially
excited about the prospect of being able to carry out
ground-breaking research into African arthropods
that conveyed disease (Report of the Government
Entomologist, 1896). In his words
To my mind the ticks present the more profitable field
for the student, whether he is interested in the systematic classification of species, in the determination
of habits and metamorphosis, in experimental
research in regard to their transmission of diseases, or
in the development of pathogenic organisms within
the body of intermediate hosts (Hooker, 1908a: 66)2.
In short, ticks were exciting. The turn of the twentieth century was an important period for the consolidation and growth in medical sciences, and the
European colonies provided plenty of opportunities
for ambitious and determined scientists to make discoveries and potentially establish an international
reputation for themselves through the expansion of
specialised journals and academic networks.
When he arrived in the Cape in 1894, Lounsbury was
well aware of work by his compatriots, Theobald
Smith and Frederick Kilborne, who in 1893 published
their conclusions that Texas fever was spread by the
bite of the popularly known cattle tick (Smith and Kilborne, 1893). So from the start he was cognisant of
the possibilities that some livestock diseases might
well be attributable to arthropods. Between 1898 and
1903 Lounsbury collaborated with veterinary
researchers and identified the primary vectors, which
transmitted all of South Africa’s economically significant tick-borne conditions: redwater, gallsickness and
East Coat fever in cattle, as well as heartwater that
can kill all domestic ruminants (Reports of the Government Entomologist, 1898, 1900, 1901, 1902,
1903, 1904a, 1904b; Lounsbury, 1900, 1904, 1906;
Norval and Horak, 2004)3. Lounsbury successfully
used international journals, such as the American
Journal of Economic Entomology, to present his discoveries to a wider audience and attain acclaim from
his peers (Hooker, 1908a, 1908b; Brown, 2003a).
2
The quotation comes from Lounsbury’s speech to the
British Association for the Advancement of Science, 1905.
3 Since 1903 entomologists have inculpated other species of
disease-bearing ticks, but Lounsbury’s original analysis
remains undisputed. See Norval and Horak for details.
307
But Lounsbury’s inspiration was not born out of scientific papers alone. As early as 1877, Cape farmers
had expressed their assumptions that ticks might be
responsible for a number of livestock diseases. In
that year, the Cape’s first state veterinarian, William
Branford, had set up a commission to hear from
farmers about their impressions of the disease situation in the colony. One of the most eloquent commentators was John Webb from the District of
Albany in the eastern Cape:
My opinion is we have a tick which made its appearance in the last 8 or 9 years. I suffered from them
then, a boutetick [sic], small like a ladybird. I was
farming on a farm without ticks, directly this tick
appeared all my stock did badly, calves died of gallsickness, boschsickness, one man lost 2 or 3,000
sheep and goats, I believe the tick caused it, I found
water on the heart, caused by inflammation brought
on by the tick. I have also shot bush bucks suffering
from the same causes, this was at Southy’s Poort, Fish
River. As this tick increases, so diseases increase, for
wherever this tick is found there are the same disease,
the tick has now travelled over 60 miles (Webb,
1877:108).
This quotation provides an insight into a number of
diseases that had not been formally identified at that
time, but were a worry to farmers who had developed their own nomenclature. Gallsickness referred
to a number of cattle diseases affecting the liver, and
by the first decade of the twentieth century had
become synonymous with anaplasmosis, conveyed
by the blue tick (Boophilus decoloratus). Farmers
also spoke of a bovine condition known as black
gallsickness, or redwater, also spread by the blue
tick and characterised by red urine. Boschsickness
was a vague term referring to any veld borne disease. By the second half of the nineteenth century,
“water on the heart” or heartwater had the biggest
economic impact on small stock populations, and
Webb was later proved correct in assuming that the
boutetick, or bont tick (Ambyloemma hebraem) was
the vector.
Webb did not refer to East Coast fever, an infection
that did not arrive in South Africa until 1902. East
Coast fever was introduced to the region with cattle
brought in from Portuguese East Africa (now
Mozambique). It proved to be the most destructive
of tick-borne diseases in South Africa, killing up to
90 percent of an infected herd. In 1903, Lounsbury
showed that it was spread by the brown tick (Rhipicephalus appendiculatus) (Lounsbury, 1904, 1906).
These early discoveries, which laid the foundations
of veterinary entomology in South Africa, revealed
how colonial science was a hybrid science. There was
no straight transfer of ideas and practices from the
imperial metropole to the colonial periphery. Veterinary entomology evolved from developments in
“germ” theory, emanating from Europe and “economic entomology”, originating in the United States
(Report of the Government Entomologist, 1896).
American research into arthropods that undermined
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the rural economy dated back to the 1870s and it
was in that context that Smith and Kilborne obtained
the funding to carry out their research into Texas
fever (Howard, 1930: 1-6; Palladino, 1996: 21-46).
In South Africa local farmers also played an important and enduring role. Even today, tick-borne diseases continue to be referred to as redwater and
heartwater, rather than Bovine babesiosis or Cowdriosis. Through observation of the veld and the parasites that occupied both the skin and the internal
organs of their livestock, farmers were able to isolate
potential vectors, even if they were unable to work
out the biological mechanisms of disease transmissions. As a result some farmers took action to deal
with ticks before Lounsbury published his entomological findings. Farmers’ experiments with chemicals and dips preceded scientific investigations into
repellents and acaricides to kill ticks (see below).
ticks were most common in the warmer, wetter,
summer rainfall areas of South Africa. These included the northern Transvaal (bordering present day
Zimbabwe and Botswana), the lowveld (which borders Mozambique and includes the area around the
Kruger National Park), all of Natal and the coastal
areas of the Cape. Ticks were less frequently
encountered on the central plateau, known as the
highveld, as they could not easily survive the cold
winters. Nor were they so common in the drier,
semi-desert areas of the western and northern Cape.
Farmers dipped their animals to protect them
against heartwater, redwater, gallsickness and East
Coast fever. Because East Coast fever was such a
devastating disease, the South African state made it
compulsory to dip animals against this infection in
the Transvaal, Natal and the eastern Cape, in order
to stop recurring outbreaks.
Historical agency can also be attributed to the environment which influenced developments in colonial
science. As the ecological studies of the 1930s and
1940s revealed, disease vectors were environmentally specific. Topography, vegetation and climate
affected the distribution of ticks, which to be pathogenic to livestock had to come into contact with a
parasite that could develop within the tick and then
be passed on to warm blooded animals. The presence of ticks was also influenced by economic contingencies. As farmers increased their flocks and
herds to meet the demands of the market, livestock
populations became denser, thereby facilitating the
spread of disease from one animal to another. Diseases that appeared to be particularly destructive
from an economic perspective attracted scientists
searching for important subjects to research, state
officials eager to ameliorate the economy and politicians seeking the rural vote. Colonial science thus
evolved from a mixture of international knowledge,
local observations in the field and environmental
and economic contingencies in a given place.
Dipping animals however was not simply about driving animals into a tank or pen for them to be treated. Dipping had scientific foundations, born out of
the laboratory as well as experiences in the field.
Once Lounsbury had identified the main vectors of
these diseases, he analysed the life-cycle of each of
the species, because that would determine how frequently animals had to be dipped. Isolating the vectors and then studying their feeding and reproductive processes was a long drawn out process, involving experiments with numerous types of ticks. This
could take months or even years to realise as there
could be seasonal fluctuations in metamorphosis
and the availability of blood meals. It also proved
difficult to keep ticks alive in the laboratory (Lounsbury, 1903). As a result entomologists like Lounsbury, became dependent on farmers, like Llewellyn
Roberts from Cottesbrook Farm in the Fort Beaufort
District of the eastern Cape, who allowed him to
carry out experiments on his property in the hope of
finding the source of heartwater, which prevailed on
his estate (Report of the Government Entomologist,
1900, 1901). Lounsbury also worked with veterinary researchers in the Cape and the Transvaal who
were particularly interesting in elucidating how ticks
acquired diseases and passed them on to livestock.
The remaining two sections of the paper will explore
veterinary entomology by looking at how scientists
and farmers tried to control these diseases, by dealing with the vector, rather than the “germ”. Tickborne conditions proved to be particularly difficult
to manage by vaccination. There was no really safe
vaccine for East Coast fever and heartwater, and
those available for redwater and gallsickness were
not immensely effective (Coetzer and Tustin, 2004).
Consequently, farmers became particularly dependent on chemical treatments, whilst ecological
analysis helped to indicate areas of potential spread.
Chemical approaches to controlling
tick-borne diseases
Following the political Union of South Africa in
1910, tick dipping became a regular part of the
farming week in areas where ticks were prevalent.
Local observations had shown that disease-bearing
So what did Lounsbury discover? In practical terms,
Lounsbury differentiated between ticks on the basis
of how many hosts they needed to fulfil their lifecycle – that is their transformations from larva to
nymph and from a nymph to an adult. Some
required one host, some two and others three,
which had implications when designing methods to
control them. Nevertheless, there were factors in
common. Female ticks laid thousands of eggs on the
ground and died after oviposition. They all required
blood for metamorphosis and reproduction and
were dependent on a certain amount of heat and
moisture, albeit to different degrees, for development to take place. Regardless of species, once
hatched from the egg, the larvae climbed to the top
of a blade of grass and awaited the passing of a
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warm-blooded host, preferably an ox. If desperate
the ticks would attack any mammal, including
humans. Finding a blood meal could take many
months and if no suitable prey appeared, the larvae
would eventually die.
In the case of the blue tick, the larva transformed
into a nymph and then into an adult on the same
beast. Given the precariousness of finding a suitable
host, their reliance on only one mammal enhanced
their chances of survival, suggesting that they had
been the most successful of the infective ticks in
adapting to their environment. Each stage of growth
was preceded by a blood meal. Once fully grown,
the female blue tick engorged for a third time and
fell to the ground to lay her eggs. The speed of this
process depended on climate, hatching and growth
being much faster in the warmer months. In the hotter parts of the country such as the Transvaal,
Lounsbury estimated that blue ticks could produce
seven generations within a year (Lounsbury, 1904,
1906). The bont and brown ticks grew in a similar
way except instead of developing on the body of a
single mammal, they dropped to the ground to
undergo the next stage of metamorphosis, and thus
needed three mammals to fulfil their life-cycle from
larval feed to egg-laying. According to Arnold Theiler, the leading bacteriologist in the Transvaal, their
chances of propagation were theoretically not so
favourable because of their need for multiple hosts
(Lounsbury, 1903; Theiler, 1904)4.
Theiler was one of South Africa’s leading researchers
and became the first Director of the Onderstepoort
Veterinary Institute, holding that position from 1908
until 1927. But there were other veterinary scientists
who also contributed to the growing knowledge of
tick-borne diseases. One such investigator was G. S.
Bruce who provided a detailed description of the
methods used in transmission experiments. Bruce
worked on East Coast fever in both Southern Rhodesia (now Zimbabwe) and Natal during the first
4 Lounsbury and Theiler used the terms African Coast Fever
and Rhodesian Tick Fever along with East Coast fever to refer
to the same disease until around 1906, when East Coast fever
became the standardised nomenclature. The variety of names
reflected on-going research into whether all these tick-borne
diseases were the same infection.
309
decade of the twentieth century. He described how
scientists collected nymphs infected with this disease
from sick cattle and placed them on the ears of
healthy bovines. Ticks were kept in place by covering
the ears with special caps that were removed when
the animal died. At death, the ticks fell into the caps
and were then kept in boxes to moult into adults.
Once fully grown, scientists placed the adult ticks
onto another bovid to see if they conveyed East Coast
fever. Proof of transmission was a rise in animal temperature within twelve to fifteen days (Bruce, 1908).
In sum, scientists showed that different species of
ticks conveyed particular disease at different stages of
their life-cycles. In the case of the one host blue tick,
the adult female picked up the redwater parasites
(Piroplasma bigeminum), or the gallsickness parasites
(Anaplasma marginale) whilst sucking on an infective
bovine, and then passed the disease onto her young
via the eggs. Blue ticks were therefore dangerous in
their larval form. The three host bont and brown ticks
were unable to hand down the disease to the next generation, but could transmit heartwater or East Coast
fever in the subsequent stage of their life-cycle. Thus
a larva that imbibed Theileria parva, the cause of East
Coast fever, or the ultravisible virus that gave rise to
heartwater (designated Rickettsia ruminantium in the
1920s) in their blood meal could pass these diseases
on as a nymph. Alternatively a nymph that became a
carrier by feeding on a sick animal could spread these
infections as an adult. Investigations showed that one
bite was enough to propagate disease and that each
type of tick discharged its pathogenic-load in a single
meal (Theiler, 1909, 1911). Consequently, these ticks
could convey a disease once in their lifetime so long
as the protozoan or micro-organism that they had
absorbed during a blood feed was one that relied on
that particular vector for its own parasitic development. The chances of a protozoan or rickettsia surviving might theoretically appear slim, but given the
extent of tick life in the summer rainfall areas, it was
relatively easy for disease to spread through a herd or
flock if the sick were not rapidly detected and isolated, and if the remaining animals were not moved to
alternative grazing land that was free of infective ticks.
Scientists also discovered that infective ticks developed at different rates and spent varying lengths of
time on the body of their hosts. This information
Table 1. An attempt to simplify the modes of transmission discussed above.
Tick
Blue Tick
(Boophilus decoloratus)
Disease
Redwater
Gallsickness
Life-cycle
1 host tick
Disease passed on by the mother through
3-4 weeks to complete its life- the eggs
cycle on a single mammal
Infective in larval stage
East Coast Fever 3 hosts tick
Brown tick
3-5 days on each host
(Rhipicephalus appendiculatus)
Bont tick
(Amblyomma hebraeum)
Heartwater
Transmission
3 hosts ticks
4-5 days on each host
Infective larva passes on ECF as a nymph
Infective nymph passes on ECF as an adult
Infective larva passes on heartwater as a
nymph
Infective nymph passes on heartwater as
an adult
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was important for devising appropriate dipping
strategies to poison specific species. The blue tick
lingered for three to four weeks on a single beast
before completing its life-cycle, hence dipping every
three weeks would be sufficient to control them.
Bont ticks tended to spend about five days on a
mammal in their larval and nymph phases and
slightly longer as an adult, whilst the brown tick
completed each stage of its life-cycle after three to
five days on each of its three hosts. Dipping to eliminate both the bont and brown tick would therefore
have to occur at least once a week, if heartwater and
East Coast fever were to be controlled (Lounsbury,
1903).
Once scientists had gathered this information, they
were now in a position to try to develop practical
measures to deal with ticks, namely through acaricides. In this respect, however, Lounsbury and his
veterinary colleagues were behind farmers in terms
of experimenting with different poisons. Since the
1890s, farmers had started to work with a range of
chemicals. These included the external application
of substances such as arsenic, which was becoming
increasingly fashionable for dealing with fruit pests,
Figure 1. Illustrations of brown ticks. The small six-legged
creature at the top is the larva. To the right of the larva is a
nymph. The 8 legged ticks at the top and in the centre are
adult males and the two at the bottom are adult females
(Lounsbury, 1904: 30).
as well as lime and sulphur solutions, commonly
used to kill the acari mites that caused sheep scab
(Brown 2003a, 2003b). Some farmers, especially in
Natal and the Transvaal, were convinced that dosing
animals with concoctions such as Stockholm tar,
garlic and aloes would cause the ticks to drop off
their hosts (Fuller, 1904; Renkles, 1908; Bedford
and Wilken-Jourdan, 1934). The evidence suggested
that even if many farmers were not suspicious of a
link between ticks and diseases, they nonetheless
wanted tick-free animals as far as possible. Ticks,
both infective and non-infective varieties, often
clung to teats, which affected milk supply, or
attached themselves to the ears and anus, causing
severe irritation or “tick worry” in an animal, which
farmers linked to decreased yields (Bedford, 1920,
1925, 1934; Story, 1920)5.
On individual farms local practices continued. But
increasingly during the first decade of the twentieth
century official scientists advocated the use of
arsenic as the most effective acaricide. How often
animals were dipped depended on which diseases
were present in a district and how long the vector
spent on an individual host. This meant every three
weeks in redwater areas and every three to five days
to kill the three host bont and brown ticks. However, arsenic was also dangerous if badly applied and
useless if too weakly administered. Some farmers
discovered that arsenic sapped the strength of oxen,
and in Natal there were complaints that dipping
reduced milk output by up to 50 percent (Theiler
and Gray, 1913).
As well as experimenting with chemicals, farmers
also tried to invent effective ways of applying solutions, by using sprays or a dipping tank. Some of the
most widely publicized accounts came from Natal.
Southern Natal, especially, had a comparatively high
rainfall and soils that farmers believed made it ideal dairy country. Ticks were prevalent in the region,
and Natal was particularly badly plagued by redwater and East Coast fever, which entered the colony
in 1906. A prominent farmer who published extensively in the Agricultural Journals was Joseph
Baynes, a prosperous cattle breeder who owned
Nel’s Rust Farm, near Richmond, Natal. He claimed
that he was the first agriculturist in South Africa to
have built a viable dipping tank on his land around
1902 (Editorial, 1902a, 1902b; Alexander, 1903).
Baynes posed as the epitome of a “progressive”
farmer, who welcomed scientific ideas and technical
innovations. He obtained his ideas for both the
tank’s design and the arsenical preparations from
Australia, where farmers were already dipping
against ticks and other vectors. This demonstrated a
trans-colonial transfer of practical farming knowledge between educated agricultural producers,
which mirrored the development of international
5
Bedford was an entomologist at the Onderstepoort Veterinary Institute.
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scientific and professional networks developed by
scientists like Lounsbury. Baynes also collaborated
with Natal’s bacteriologist, Herbert Watkins-Pitchford, and together they worked on developing a safe
and effective dip that could be used as regularly as
three days to deal with the brown tick. Initially, they
tested a range of proprietary dips that were appearing on the South African market and found that
some were abrasive on the skin whilst others were
downright dangerous, leading to arsenic poisoning.
After a large number of trials, Baynes and WatkinsPitchford’s came up with a formula, unimaginatively named “laboratory dip”. This consisted of two
pounds of arsenite of soda to 100 gallons of water.
Due to the need to dip frequently in the case of East
Coast fever, the strength of the arsenic was less than
that used to combat the blue tick. They also added
soap and paraffin to reduce scalding and to deposit
a residue on the skin that increased the killing power of this toxicant (Watkins-Pitchford, 1909, 1914;
Editorial, 1913).
The Veterinary Department recommended the use
of ‘laboratory dip’ in the East Coast fever areas after
1910, although they later extended the interval
between immersions from every three days to a
more viable five day routine, given that the brown
tick could spend up to that length of time on a single beast (du Toit and Viljoen, 1929). Researchers
such as Arnold Theiler believed that they could wipe
out East Coast fever because it was a recently
imported disease and had not spread throughout the
country, so long as “the dipping has been carried out
energetically enough and for a sufficient length of
time before the disease has been introduced on a
farm” (Theiler, 1911: 505). The idea was that if livestock had been properly cleansed over several
months, the tick population would have declined to
such a degree that there would be too few arthropods on the land to enable an epizootic to break out
on a farm. In effect the cattle acted as bait as they
attracted the ticks that were then annihilated in the
dipping tank. To encourage farmers to erect tanks,
which were expensive, the government introduced
legislation in 1911 facilitating the procurement of
loans from the Land Bank, repayable over ten years
at low rates of interest. The Native Affairs Department also obtained advances to construct tanks in
the African reserves and recouped the cost from
African men who were liable for a 5s dipping tax
(Anon, 1911; Dower, 1912).
To qualify for these funds, farmers and administrators had to build tanks according to veterinary recommendations, which appeared in various official
publications such as the Agricultural Journal. The
favoured design was the plunge tank because the
depth ensured complete coverage with the acaricide.
Herders drove cattle from a crush pen down a
fenced race, which dropped into a five to six foot
dipping tank, through which the animals swam and
then staggered into a yard to dry off. The tank was
311
sunk into the ground, lined with concrete and supported with iron rods. In Natal many farmers located their tanks near the homestead where the water
supply was most copious. Some farmers built dams
to conserve water specifically for the dipping tank
(Theiler and Gray, 1912; Cleghorne, 1914; Webb,
1920). Because dipping against East Coast fever was
so regular, other ticks would be killed also. There
were no compulsory dipping orders in areas free of
East Coast fever, but some stockowners outside the
quarantines zones erected tanks to eliminate endemic redwater and heartwater (Theiler and Gray, 1912,
1913; Theiler, 1921).
Figure 2. Picture showing a bovine swimming through the
dipping tank on Mr McDougall’s Farm, East London District,
eastern Cape. The tank was 71 feet long and 6.5 feet deep.
It held 6,360 gallons of solution (Editorial, 1906).
Dipping initially proved to be relatively effective, but
it was also a costly and time consuming occupation.
Because many species of tick, including the brown
tick, settled in the ear canal or around the anus –
areas that tended to be insufficiently saturated in the
tank – stockowners also had to hand-dress their animals. Scientists recommended a variety of mixtures
for this including paraffin oil and Stockholm tar
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preparations, as well as ordinary dipping solution, all
of which were applied with a swab (Dixon, 1911;
Watkins-Pitchford, 1914; Webb, 1920). Farmers
together with the stock inspectors, who were
employed by the state to monitor the dipping procedures in the African reserves as well as on settler
farms, also had to make sure that the arsenic solution remained at a particular strength. Rain water
diluted the solution in the tank, whilst hot dry
weather caused liquid to evaporate, concentrating
the chemicals and increasing the likelihood of skin
damage. Watkins-Pitchford developed a tool known
as an isometer to measure the strength of the dip
(Watkins-Pitchford, 1914). Nevertheless the expense
involved in procuring the chemicals, as well as the
effort required to mix safe solutions, which was particularly difficult in times of drought when water for
the tanks was scarce, could reduce the frequency and
the efficiency of the dipping operations.
Figure 3. Watkins-Pitchford’s Isometer (Watkins-Pitchford,
1914: 122).
The most irksome regulations and procedures surrounded East Coast fever. Government stock inspectors oversaw operations in the African reserves and
periodically checked for enforcement on settler
farms also. There were high fines for failing to dip
regularly and concealing cattle. The greatest hardships occurred in the African reserves, where people
had to travel long distances to a dipping tank. Traditionally, men looked after cattle, but as more and
more males sought paid work in the cities and mines,
the burden of managing bovids fell onto women. In
Natal, which proved the hardest region to clear of
the disease, Zulu women objected to having to trek
several miles, twice a week to the communal dipping
tank and then loiter around whilst hundreds of cattle waited their turn. Women also complained about
having to hand-dress the ears of stock, as this was
particularly onerous. In addition, some animals occasionally drowned in the dipping tank and there were
concerns that the arsenic solutions scalded the skin,
because they were not properly prepared (Inspector
of Locations Estcourt, 1920; Resident Magistrate
Pinetown, 1923; Wheelwright, 1923; Resident Magistrate Nongoma, 1925). Some white farmers also
remonstrated against the process, abhorring the cost
and effort, and dubious about the effectiveness of
dipping, given that tick-borne diseases remained a
major problem facing pastoral producers (Dicke,
1920; Gray, 1920; Viljoen, 1920; Editorial, 1926).
Nevertheless, despite the difficulties and objections,
dipping was relatively effective at controlling, but
not eradicating, specific diseases. By the second
decade of the twentieth century, all the most dangerous tick-borne diseases were in decline and gradually more and more farmers invested in tanks. By
the 1920s Natal possessed the greatest number of
dipping facilities with an average of one tank per
300 cattle. Elsewhere, where occurrences of East
Coast fever had become far rarer, the figure was
nearer one tank per 1000 animals, with several hundred beasts passing through them each day (except
Sundays) (Lawrence, 1992). Richer farmers, who
produced livestock for a commercial market had
their own tanks, but communal dipping was the
norm in the African reserves. Significantly, dipping
was instrumental in enabling the eradication of East
Coast fever from South Africa. In 1954, the Director of Veterinary Services, Raymond Alexander,
announced that the disease had disappeared as a
result of dipping, strict quarantines in affected areas
and the slaughtering of infected herds to destroy any
carrier animals. Over the previous 50 years, East
Coast fever had claimed the lives of approximately
5,500,000 cattle in Natal, the Transvaal and the
eastern Cape (Diesel, 1948, 1949; Alexander,
1955). The eradication of East Coast fever enabled
farmers to increase their herds, and also eased the
work load on farms, as stockowners were relieved of
the burden of mandatory cattle dipping. Although
dipping against East Coast fever had been difficult,
and at times unpopular, its ultimate elimination
showed that dipping could be an effective weapon
against tick-borne diseases. However, the fact that
dipping alone did not lead to its disappearance, indicated that there were limitations to this strategy as
it was impossible to destroy all the ticks. Dealing
with East Coast fever demonstrated that a variety of
methods were needed to mitigate the impact of tickborne diseases.
The need for a variety of initiatives became particularly clear in relation to blue ticks. Optimism about
the viability of dipping over the longer term came
under threat in 1938 when farmers from the East
London District in the eastern Cape complained that
arsenic no longer destroyed all the blue ticks and
their cattle were succumbing to redwater. Initially
they blamed the state for allowing retailers to sell
sub-standard chemicals, but it soon became clear
that the quality of the poison was not the problem.
Research at Onderstepoort suggested that some blue
ticks had evolved a resistance to the dips (du Toit et
al., 1941; du Toit, 1943). The chemist Graeme White-
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head attributed this to changes in the enzymatic
structure of the outer shell which created a barrier
against the absorption of the toxins (Whitehead,
1975)6. Resistance to chemicals was nothing new
from a scientific perspective, as American fruit and
potato growers had observed a decline in the effectiveness of crop spraying with the usual compounds
in the 1910s. Now the problem had spread to the
livestock industry (Whitenall and Bradford, 1945).
Stockbreeders in countries such as Australia,
Uruguay and Nyasaland (now Malawi) also discovered that ticks were starting to counteract the chemical controls and this generated correspondence
between Onderstepoort and scientists elsewhere
(Alexander, 1941, 1951; du Toit, 1942). The experience of chemical resistance was thus becoming a
global problem and encouraged an exchange of
knowledge not only with scientists working in the
British colonies, but also with researchers in other
parts of the world. Genetic adaptation of any one
species of tick might initially only affect a small area,
as was the case with the blue tick in East London,
but there was always the possibility that this problem might spread not only within a country but also
across borders through trade. Mutual interest therefore fostered collaboration.
Despite blue tick resistance to arsenicals, South
African entomologists and vets continued to place
their hopes in a chemical solution to the redwater
problem. Following the Second World War, the possibilities of destroying ticks with the new synthetic
chlorinated hydrocarbon insecticides such as DDT
and BHC became apparent (Palladino, 1996: 2628)7. Scientists at Onderstepoort began experiments
to test the efficacy of DDT and BHC on a number
of insects and parasites, including mosquitoes, tsetse
flies and ticks. The leading researcher into ticks was
Gertrud Theiler, daughter of the first Director of
Onderstepoort (Arnold Theiler) who also organised
the ecological survey of their distribution (see
below). One of the few female scientists at Onderstepoort at that time, Gertrud Theiler worked with
the field vet, Alexander Diesel, to try to come up
with an acaricide that was effective against a range
of ticks. Now that the blue tick proved to be developing a resistance to arsenicals, there was evidently
the fear that the brown and bont tick would follow
suit. Their work united the laboratory and the field
as experiments involved liaising with farmers in the
East London District who provided the scientists
with livestock and land on which to carry out their
trials. Experiments demonstrated that DDT and
6 Whitehead worked in the Research Department of the
commercial company, African Explosives and Chemical Industries Ltd, showing liaison between the state and industry in
developing chemical products.
7 The Swiss Company, Geigy, invented the synthetic insecticide DDT in the late 1930s and it was first used agriculturally in the United States to destroy the Colorado Potato Beetle.
313
BHC killed the blue tick (Theiler, 1944; Diesel,
1947; Whitenall and Bradford, 1947; Fiedler, 1952).
Nevertheless, once again chemicals failed to provide
the ultimate solution. Many farmers could not
afford to use these costly preparations and it was
difficult to keep them at a standard and effective
strength. Those who did try the new acaricides used
them to protect animals against heartwater and East
Coast fever too. However, in 1949 Theiler reported
that some eastern Cape farmers had experienced an
increase in bont and brown tick populations on their
animals because these compounds were not so effective against these varieties of arthropods. Theiler
saw this as a disaster because the three host ticks
had almost been dipped out of the eastern Cape by
the late-1940s and heartwater and East Coast fever
were no longer the cause of high mortality in this
region (Theiler, 1949a). Now there was no universal
acaricide capable of destroying all types of infective
ticks. This eventuality threatened to increase the
expense of “modern” pastoral farming and presented problems in terms of livestock management and
safety if animals had to be immersed more frequently in different solutions. Theiler proposed further research into these chemicals. But by 1950
some arsenic resistant blue ticks had ceased to be
affected by DDT and BHC too (Whitehead, 1956).
The blue tick, therefore, was particularly adept at
developing a resistance to chemical dips.
This outcome presented a crisis for entomologists and
tested the credibility of their scientific practices. It led
not only to more research into dips, but also encouraged a reappraisal of the inter-relationship between
livestock and disease from the 1950s. Some scientists, including Arnold Theiler back in the early twentieth century, had argued that dipping alone might
not eliminate well-established endemic diseases and
that animals should be exposed to ticks whilst young
in order to build up some natural resistance to infections such as redwater, which did not have the same
mortality toll as East Coast fever (Theiler, 1905). This
idea was based on the observation that imported cattle were particularly susceptible to tick-borne diseases, suggesting that “indigenous” livestock, or at
least lineages that had lived in a particular South
African environment for a long time, had developed
some tolerance to redwater and heartwater. However,
many farmers were reluctant to expose their valuable
animals to tick strike, not only because of a lack of
vaccines, but also due to a lack of cures. It was only
in the 1950s that scientists could advocate reasonably
effective treatments for both redwater and heartwater
(Henning, 1949). By that time East Coast fever had
disappeared from South Africa, so the availability of
drug treatments made it more feasible for farmers to
expose their animals to a certain amount of tick activity in endemic areas.
In addition, researchers and farmers also carried out
breeding experiments to try to produce types of live-
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stock that were more resistant to infections. Indigenous varieties of livestock tended to be small, presenting low milk and meat yields, hence encouraging commercial farmers to import pedigree stock
from Europe, which were more vulnerable to disease. From the 1930s the Agriculture Department
sponsored research into the cross-breeding of local
with exotic cattle at the experiments stations at
Mara and Messina in the northern Transvaal (Agriculture Department, 1934). These experiments ultimately produced the Bonsmara breed of cattle that
was less susceptible to protozoal infections. Drugs
and environmentally tailored breeds therefore
reduced the dependence on regular dipping, and following the disappearance of East Coast fever in the
1950s, enabled farmers to engage in a wider range
of measures to tackle tick-borne diseases (Interviews, 2003). This approach was not only contingent upon advances in science, but also on a clearer understanding of the environment ticks inhabited
and their biological interaction with other species.
This information emanated from Gertrud Theiler’s
zoological survey into ticks.
Ecological Entomology:
South Africa’s first tick survey
The Zoological Survey (1936-1944), reflected a
growing interest in ecology, which some historians
have already reflected upon in the context of the
British Empire (Kolbe, 1982). Ecology was an
emerging intellectual field during the inter-war years
that involved an inter-disciplinary approach to
understanding the interrelationship between the animal and plant kingdoms. The underlying agenda was
to further sustainable agricultural growth based on a
thorough realisation of the constraints and opportunities presented by a given environment. Developments in ecological thinking emanated not only from
Europe and its colonies but also from the United
States. In 1920 the American Ecological Society
began to produce its journal, Ecology, which became
an important forum for intellectual exchange and
contributed to the international spread of scientific
ideas about the natural world (Robin, 1997; Anker,
2001; Tilley, 2001, 2003). During this period, medical ecology also became a key element in epidemiological research as investigators tried to ascertain the
provenance of new diseases and the reasons why
they could assume epidemic proportions (Mendelsohn, 1998). In the context of entomology and veterinary medicine in South Africa, ecological as
opposed to purely biomedical approaches to understanding the distribution and source of diseases
focussed on studies into nagana (bovine trypanosomosis) and ticks. Botanists, meanwhile, carried out
studies of soils and flora in order to understand the
interrelationship between topography, climate, soils
and vegetation (Bews, 1916, 1925; Pole Evans,
1922; Acocks, 1953). Types of vegetation affected
the habitat and thus the dispersal of arthropods.
Gertrud Theilers tick survey highlighted the linkages
between disease and environment and resonated
with Libby Robin’s notion that ecology was a “science of empire”, and an epistemological instrument
for environmental exploitation (Robin, 1997). The
assumption was that if scientists and farmers understood the local ecology, they could take steps to alter
the environment or the management of farms in
order to reduce the chance of disease. Scientists thus
regarded ecology as a tool of development.
Theiler believed that mapping out the distribution of
ticks, noting their preferred habitats as well as the
influence of climate on seasonal changes in reproductive rates was an important tool for giving scientists a better understanding of the environment
that sustained tick life. Theoretically, this would
help entomologists to demarcate the areas to which
vector-borne diseases might propagate, so that farmers could be warned of the need to take action to
protect their stock. It could also reveal species density in a given area and thus the probability of disease. In her resumés of the findings, published in
the Onderstepoort Journal between 1948 and 1953,
Theiler argued that
In spite of the shortcomings inherent in such a generalized survey, it has nevertheless been possible in
many instances to draw definite conclusions as to the
factors encouraging or discouraging the increase of
various ticks species, and hence to their distribution
in South Africa (Theiler, 1948: 218).
In order to collate this information, field vets were
asked to divide their districts into three or four
zones, each reflecting variations in altitude, vegetation and climatic patterns. Within each of these
areas, they chose about four farms from which to
collect ticks, at different times of the year, from a
variety of domestic animals. Farmers also contributed and helped to identify prevalent species on
their land. Participants placed selected parasites into
test tubes and sent them to Onderstepoort, along
with a description of the climate and vegetation in
the region from which they came. Once again, entomological work into ticks required close interaction
between events in the field and processes in the laboratory.
In many respects the results that Theiler collated
between 1937 and 1948 did not live up to the ideal,
and formed a somewhat rough and ready picture of
tick distribution on the ground. In her overview of
this enterprise Theiler, rather ungraciously, complained that many of the farmers and veterinary surgeons who volunteered to participate in the survey
had little understanding of ecology. She claimed the
majority were poor at identifying and selecting the
variety of specimens needed to form a representative
sample of national tick life. In her words “few persons have a species-sense and too frequently the identification is not only inadequate but also inaccurate;
and in any case the popular name, in most instances,
is generic rather than specific” (Theiler, 1948: 218).
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Theiler also found her collectors erratic in their
efforts, making it difficult to get a clear comparative
picture of seasonal and annual variations in tick populations, as well as their distribution patterns, during
the eleven years of this survey. The study also failed
to provide adequate insights into the rate of tick
growth and propagation in the field, as entomologists
could only really observe these processes in the laboratory. Laboratory techniques helped scientists to estimate reproductive as well as developmental and survival periods of individual species, but because it was
impossible to replicate the daily changes in temperature and moisture levels of an outdoor environment,
their conclusions were an approximate, rather than
an accurate, indication of tick behaviour in the veld
(Theiler, 1948). There were thus inadequacies in the
methodology and findings from both the laboratory
and the field.
On the positive side, Gertrud Theiler did claim that
the investigation confirmed some long-held suppositions. It revealed that although many of the dangerous
species inhabited the summer rainfall areas, there
were nonetheless some marked differences in environmental preferences that helped to explain why
more or less of a particular tick might be found in a
specific locality. The survey affirmed that the main
limiting factor in their distribution was climate. This,
along with soil types influenced the nature of the vegetation, which might or might not attract tick life. The
evidence corroborated earlier suspicions that the blue
tick was the most environmentally adaptive of the
infective varieties. It could survive almost anywhere
apart from the arid districts of the Cape. The absence
of rainfall was therefore the key constraint on its
spread. In terms of vegetation, the blue tick was versatile, enjoying forested river valleys as much as open
grasslands (Theiler, 1949b). The fact that the blue tick
needed only one host to complete its life-cycle, as well
as its ability to survive in all but the driest of environments, favoured its chances of propagation. The
blue tick had the widest distribution of the three
species. Hence cattle were more likely to contract redwater or gallsickness than they were heartwater or
East Coast fever.
The bont tick, on the other hand, was far more environmentally localised. Compared with the blue tick,
precipitation was less of a limiting factor and the
main direct check on its distribution was the type of
vegetation. In the Transvaal and Natal the bont tick
favoured bushes and thickets, whilst in the grassy
eastern Cape it was largely restricted to the river valleys. Like tsetse flies, the bont tick required shade to
avoid desiccation and it was unable to survive in
areas devoid of trees or shrubs. This arthropod was
most active in the summer, hence heartwater tended
to break out in the hotter months (Theiler, 1948).
Finally, the brown tick resembled the blue in that a
lack of rainfall constrained its proliferation. But it was
more akin to the bont tick in terms of its vegetative
predilections. The brown tick was absent from arid
315
districts as well as the highveld where the winter
frosts jeopardised its survival. By the 1940s the
brown tick appeared to have had a relatively limited
distribution in the heavily populated African reserves
in the eastern Cape. Theiler attributed this to overgrazing, emphasising the importance of grass and
shrub cover for the survival of this species (Theiler,
1949c). However, entomologists and vets did not recommend keeping too many animals on a farm as overgrazing could destroy the grasslands. Overgrazing was
a major problem in some parts of South Africa, especially in the African reserves. Although state scientists
viewed overstocking as a threat to pastoral sustainability, ironically, intensive grazing might have contributed to the elimination of East Coast fever from
the eastern Cape by removing vital vegetation.
Overall, the survey provided a generalised impression of vegetation preferences and gave some hints
as to the effects of climatic factors on tick life. However, many questions remained unanswered. Writing
ten years after the study, Theiler argued that scientist
were still unsure as to how seasonal variations in
temperature and moisture levels really affected the
development and activity of these parasites.
Undoubtedly tick reproduction was faster in the
warmer and wetter summer months, but a more
nuanced understanding of this interrelationship was
needed to predict prevalence. The blue tick appeared
to be active all year round, but the influence of
weather patterns on the annual distribution of bont
and brown ticks, which were more sensitive to floral
changes, remained unclear. The survey also looked at
the effect of veld burning on tick survival. Again the
results were inconclusive. Many African and settler
farmers had traditionally burnt the grasslands, not
only to speed up the germination of fresh vegetation,
but also to reduce the number of ticks. But whether
the heat penetrated deep enough into the ground to
destroy a significant number of eggs remained scientifically unsubstantiated (Theiler, 1959).
This ecological survey thus drew upon various fields
of knowledge. On the one-hand, Theiler and her colleagues looked to international science to suggest
methods of examining the nature of tick infested
environments. On the other hand there was also
considerable local input, highlighting the importance of specific localities in influencing the distribution of particular types of ticks and specific diseases. Farmers might not have had the most precise
taxonomical knowledge, but they played a key role
in collecting specimens that fed on their animals,
thus furthering research into South Africa’s parasites. The survey also showed that farmers continued to pursue older methods of disease control,
such as veld burning, alongside the introduction of
chemical treatments. This indicated that stockowners did not have total faith in the efficacy of chemicals. Growing tick resistance to dips suggested that
this was probably a wise approach. Ultimately, in
the absence of legislation for administering gallsick-
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ness, redwater and heartwater, farmers were left to
their own devices and tended to engage in practices
that appeared to work. Knowledge became
hybridised as dipping existed alongside not only veld
burning, but also the dosing of animals with a range
of concoctions such as garlic and aloes, as discussed
in relation to East Coast fever earlier in this paper
(Bedford and Wilken-Jourdan, 1934).
Whether veld burning reduced tick presence was a
debatable point, as was the role wildlife might play
in spreading these diseases. By the mid-twentieth
century, scientists knew that game constituted the
reservoir of a number of livestock infections, most
notably bovine trypanosomosis. During the 1930s
and 1940, Wilhelm Neitz, a parasitologist based at
Onderstepoort, carried out a number of experiments
to ascertain whether various species of antelope
could be the carriers of tick-borne diseases. He took
blood from game animals, which demonstrated no
symptoms of disease, and injected it into livestock
to see if the latter fell sick. He also fed ticks on
game animals then transferred the arthropods to
cattle and sheep. In this way he discovered that the
blesbuck (a sort of antelope) and the black wildebeest harboured the rickettsia that caused heartwater, and speculated that springbuck might also be
carriers (Neitz, 1935, 1937, 1944). Consequently
heartwater could survive in nature, regardless of the
presence or absence of domestic animals. Given that
heartwater was endemic in South Africa, it was possible that some types of game had been the initial
source of the disease in domestic livestock. Neitz
also noted that in the wild, bont ticks readily fed on
antelopes, strengthening the possibility that game
might be a perpetual source of infection (Neitz,
1937). This linked in with earlier observations of
farmers, like John Webb, who had associated ticks
with wildlife back in the 1870s.
The presence of the heartwater microbe in game thus
illustrated the complexity of the relationship between
livestock, ticks, disease, vegetation and wildlife and
raised questions as to how tick-borne diseases could
be controlled in a country where game were abundant. Since the 1880s, South African governments
had started protecting wildlife through regulating
hunting and establishing game reserves. The conservationist and sporting lobbies were politically influential and obtained state support for the protection of
wildlife for tourism, licensed hunting and scientific
research. By the 1930s a number of large wildlife
sanctuaries existed, including the Kruger National
Park in the Transvaal and the Umfolozi-Hluhluwe
Game Reserve in Natal – both areas of high tick density. Keeping wildlife off farm land was not always
possible as the game reserves were often unenclosed,
so it was easy for ungulates to roam onto adjoining
properties. Thus the survival of wildlife was epidemiologically problematic (Mackenzie, 1988; Carruthers,
1995; Brooks, 2001; Brown, 2002, 2008b). Neitz
argued that “it is quite impossible to eradicate the
tick-borne diseases of domestic animals from an area
where game and their movements are not as vigorously controlled as all other animals” (Neitz, 1944:
26). The only solution was to fence off game reserves
and farmlands effectively, thereby confining wildlife
to designated areas. This information served as a
warning to stockowners to fence their properties and
try to keep livestock away from pastures frequented
by wandering game.
Conclusion
Ticks were a major threat to the South African economy and from the late nineteenth century they were
subjected to extensive entomological and veterinary
research. Although South Africa was not the first
country to discover a link between ticks and disease,
the work of Lounsbury in particular was pioneering
in the African context, and he and his colleagues
revealed the mode of transmission of some of the
country’s most dangerous infections. By the 1930s
scientists began to look at the geographical distribution, as well as the environmental constraints on tick
spread in order to ascertain areas that were potentially vulnerable to some form of arthropod-borne
epizootic. There were too many ticks to make eradication of these parasites feasible, but scientists did
devise dipping techniques to try to reduce endemic
diseases and used this method along with quarantines
and slaughter policies to tackle East Coast fever.
However, dipping soon revealed its limitations. As the
blue tick became resistant to arsenic and later to the
BHC and DDT acaricides, scientists looked for new
ways of managing parasitic diseases. Scientists, therefore, had to continually adapt their research and control strategies to meet the challenges of a changing
entomological and epidemiological landscape.
The paper also aimed to show that farmers were at
the forefront of entomological observations and they
were ahead of scientists at identifying ticks as potential propagators of disease, as the evidence of John
Webb before the 1877 Stock Commission demonstrated. Farmers also took the lead in devising dips
and tanks to try to kill ticks they believed harmed
their animals. Lounsbury learnt much from farmers’
knowledge, carrying out experiments on vectors they
assumed transmitted diseases, such as the bont tick.
Lounsbury was also influenced by work in the United States that had confirmed ticks spread Texas
fever. Australian farmers, who were already dipping
their stock to counteract a range of arthropods, were
inspirational from a practical sense as their published accounts encouraged South African farmers
like Joseph Baynes to carry out dipping experiments
on their own properties. Entomological science as it
evolved in the British colony of South Africa thus
drew upon a number of pools of knowledge, which
included farmers’ experiences in the field as well as
scientific experimentation in the laboratory. Gertrud
Theiler’s tick survey, as well as Neitz’s zoological
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research into linkages between tick-borne diseases
and wildlife showed that South Africa had also
bought into the relatively new, but rapidly expanding, field of ecology. The epidemiological mapping
that this entailed, revealed that tick-borne diseases
would not be easily eradicable due to their widespread distribution and the fact that wildlife constituted not only a reservoir of heartwater (and possible other tick-borne diseases) but also hosts for ticks.
Wildlife thus functioned as both a potential reservoir
of disease and a carrier of infective ticks.
The widespread distribution of tick-borne diseases,
as well as limitations to dipping, as first revealed in
relation to redwater, led to a revision in thinking
about the control of these infections, which pervades
to this day. In the modern parlance, the recommendation now is for farmers to adopt an “integrated
approach”. During the second half of the twentieth
century, scientists have advised a mixture of measures, including exposure to ticks in endemic areas
as well as breeding more disease-resistant types of
livestock, such as the Bonsmara cattle. Vaccinations
do exist, but these are attenuated live vaccines,
which are hard to store and not terribly safe, often
having to be accompanied by antibiotic treatments to
forestall serious reactions. Many farmers in South
Africa cannot afford to invest in such precarious
treatments. As part of this “integrated approach” to
disease control, dipping is now just one of several
strategies endorsed to mitigate the impact of these
vectors. This shift in thinking only became possible
after the eradication of East Coast fever in the early
1950s. Redwater, gallsickness and heartwater do not
normally cause as high mortality as East Coast fever
and many animals recover preventing the need for
the strict regulations that characterised the campaign
against East Coast fever. Nonetheless, there is always
the danger that East Coast fever could be re-introduced and new diseases are constantly emerging.
South Africa’s tick environment will continue to provide challenges and opportunities for entomologists,
parasitologists and farmers alike (Interviews, 2003).
Acknowledgements
I would like to thank Heloise Heyne, Arthur Spickett and Mick
Combrink from the Department of Parasitology at the Onderstepoort Veterinary Institute, for sharing their knowledge and
experience of tick-borne diseases with me. I am also grateful for
the advice and support received from Rudolph Bigalke, Theuns
Naudé and David Swanepoel, also from the Onderstepoort Veterinary Institute. William Beinart and Daniel Gilfoyle provided
useful comments from a historical perspective. Finally, I would
like to thank Anne Hardy, Annick Opinel and Gabriel Gachelin for
organising the medical entomology workshops at which this
paper was presented, as well as the Wellcome Trust for funding
my post doctoral study of South African veterinary medicine.
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Sowing the seeds of Economic Entomology:
houseflies and the emergence of Medical Entomology in Britain
J.F.M. Clark
Institute for Environmental History, University of St Andrews, St Andrews, Scotland, UK.
Abstract. The golden age of medical entomology, 1870-1920, is often celebrated for the elucidation of the
aetiology of vector-borne diseases within the rubric of the emergent discipline of tropical medicine. Within these triumphal accounts, the origins of vector control science and technology remain curiously underexplored; yet vector control and eradication constituted the basis of the entomologists’ expertise within
the emergent specialism of medical entomology. New imperial historians have been sensitive to the ideological implications of vector control policies in the colonies and protectorates, but the reciprocal transfer of vector-control knowledge, practices and policies between periphery and core have received little
attention. This paper argues that medical entomology arose in Britain as an amalgam of tropical medicine
and agricultural entomology under the umbrella of “economic entomology”. An examination of early twentieth-century anti-housefly campaigns sheds light on the relative importance of medical entomology as an
imperial science for the careers, practices, and policies of economic entomologists working in Britain.
Moreover, their sensitivity to vector ecology provides insight into late nineteenth- and early twentieth-century urban environments and environmental conditions of front-line war.
Key words: history of entomology, medical entomology, economic entomology, housefly, infant welfare,
war.
The history of medical entomology has often been
subsumed within that of tropical medicine. Consequently, triumphal accounts of the elucidation of the
aetiology of vector-borne parasitic diseases between
1870 and 1920 abound (Philip and Rozeboom,
1973; Service, 1978; Bruce-Chwatt, 1988; Busvine,
1993). With the socio-cultural turn in the history of
science and medicine, some historians have emphasized the medical and social contingencies that lay
behind the creation of institutions; and they have
sought to retrieve the social and economic circumstances within colonies and protectorates that were
often masked by the ideological programmes that
inspired the “universal” specialism of tropical medicine.
Both triumphal and more socially and politically
sensitive histories, however, provide similar accounts
of the development of the specialism of tropical medicine: a golden age of the aetiology of vector-borne
diseases was followed by metropolitan-based “vertical” campaigns of vector eradication or prevention
(Worboys, 1993). The close link between parasitevector diseases and the Mansonian vision for tropical
medicine has meant that the history of medical entomology has often been indistinguishable from tropical
medicine. The ideology of “constructive imperialism”
undoubtedly played a central role in the institutionalization of medical entomology in Britain, but tropical
medicine’s commitment to vector control drew upon
established knowledge and institutions within agricultural science.
Correspondence: J.F.M. Clark, Institute of Environmenal History, University of St Andrews, St Andrews, KY16 9AL, Scotland, UK, e-mail: jfc2@st-andrews.ac.uk
To locate the discipline within the specialism of
tropical medicine, progressive celebratory histories
of medical entomology recount early speculations
about the connections between insects and disease
(Nuttall, 1900; Service, 1978: 603-608). More
appropriately, the early history of medical entomology should be located in the development of a specialism of applied, or “economic”, entomology,
which sought to identify and control insects that
were baneful and beneficial to agriculture and
human health. Consequently, early medical entomologists were less intent than their counterparts in
tropical medicine on establishing a knowledge base
that was unique to the colonies and protectorates.
As entomologists, they actively sought to translate
their colonial experiences to the metropolitan context. Whereas Patrick Manson excluded fly-borne
enteric diseases from his definition of tropical medicine (Worboys, 1993: 520), entomologists played a
key role in the “fly danger” campaigns that occupied
Britain and other temperate lands in the early twentieth century: they argued that flies, as mechanical
vectors of infantile diarrhoea, were to Britain what
mosquitoes were to the tropics (Anonymous, 1907).
Anti-housefly campaigns were, therefore, a concomitant part of the emergence of medical entomology in Britain.
Experience in the British colonies permitted emergent entomological experts to redeploy their expertise in agricultural entomology in the cause of medical entomology. As an institution or discipline,
applied entomology in Britain was forged from agricultural science and tropical medicine, under the
umbrella term of economic entomology. Although
significant, colonial experiences of crop pests and
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J.F.M. Clark - Medical Entomology in Britain
vectors of disease were just one facet of the complex
interplay of humanity and insects that shaped the
development of medical entomology. A convergence
of environmental factors in Britain led to a significant increase in houseflies and a concomitant rise in
infant mortality in the closing decades of the nineteenth century. Changes in agriculture, war-time
experiences, increasing urbanization, urban and
suburban growth, and a “refuse revolution” all
helped to shape the interplay of economic entomologists and insects in Britain. This paper argues that
the house-fly scare constituted a significant facet of
what Timothy Mitchell has called “the rule of
experts” (Mitchell, 2002).
Economic Entomology
Economic entomology traditionally occupied the
borderlands of agricultural and natural science.
Throughout the last several decades of the nineteenth century, there was a small group of agricultural entomologists working in Britain; they had little in common with the priorities of the metropolitan entomological community that coalesced around
entomological societies and museums. Instead, agricultural and horticultural societies, new agricultural
colleges, and county extension schemes provided
them with support and legitimacy. Operating on the
periphery of entomological science, they gained new
status when Joseph Chamberlain’s “constructive
imperialism” called upon applied science to exploit
Britain’s colonial possessions. The identification and
eradication of insect pests assumed a prominent role
in Chamberlain’s imperial designs (Clark, 2001).
Nevertheless, entomologists struggled to define a
clear role for themselves in this imperial programme. In the rapidly emerging field of medical
entomology, practitioners of biomedicine, who were
often untrained in entomology, studied the aetiology
of insect-borne diseases. Some entomologists highlighted the importance of taxonomy for the correct
identification of economically harmful or beneficial
insects. Classification work was, however, principally based in metropolitan museums, and was perceived as the preserve of “pure” systematists. In
contrast, the most vociferous exponents for a profession of applied entomology linked their fortunes
to the development of expertise in agricultural entomology. Robert Newstead (1859-1947) and F.V.
Theobald (1868-1930), for instance, demonstrated
the continuities between these agricultural and medical applications.
In 1899, E. Ray Lankester, Director of the British
Museum (Natural History), retained the services of
F.V. Theobald, entomologist at the South-Eastern
Agricultural College, to identify mosquitoes from
tropical and sub-tropical colonies. Between 1901
and 1907, Theobald received 21,200 specimens,
and purported to name 275 new species (Theobald,
1901-1907). Theobald’s early taxonomic study of
British Diptera justified his involvement in imperial
entomology. His established reputation as an economic entomologist was also a contributing factor
to his employment with the museum. This dual role
captured the ambivalent position of the metropolitan-based economic entomologist. At the same time
that Theobald provided purely systematic work on
the mosquitoes of empire, he assisted the Director
of the museum with reports on economic zoology
for the Board of Agriculture (Theobald, 1903).
In Britain, agricultural entomologists provided the
necessary experience and expertise in insecticide
applications. In April 1911, Robert Newstead –
elected in 1905 as lecturer in economic entomology
and parasitology – became the first Dutton Memorial Professor of Entomology at the Liverpool School
of Tropical Medicine (established 1899). Prior to
assuming his position at Liverpool, Newstead had
devoted his entomological investigations exclusively
to agricultural applications. He compiled the general index to Eleanor Ormerod’s Annual Reports of
Observations of Injurious Insects, 1877-1898
(1899) after declaring a desire to emulate her entomological career (Laing, 1947; Anonymous, 1947).
Newstead became a specialist in the study of scale
insects (Newstead, 1901-1903). This subject attracted considerable attention when San José scale
threatened to destroy the Californian citrus fruit
industry in the 1880s. D.W. Coquillet’s successful
eradication of the pest with a new fumigant – hydrocyanic-acid gas – was proclaimed as one of “the
great achievements in the entomological history of
California” (Rothman, 1987: 226-229). Newstead
made numerous experiments with the American
insecticidal technology on scale insects in Britain.
When he shifted his emphasis to medical entomology, he applied this same insecticidal technology. In
1915, he employed hydrocyanic-acid gas against bed
bugs (Newstead, 1899-1900, 1902).
Economic entomology achieved respectability
between 1890 and 1914 through the creation of
specialist educational programmes and acknowledged careers in the field. Imperial ambitions acted
as a catalyst for this process, but the intellectual
foundations of the discipline were laid earlier. The
British metropolitan entomological community,
which had long spurned economic entomology,
increasingly embraced this specialist facet of their
discipline. Although metropolitan museums continued to be centres of “pure” insect systematics, they
attracted personnel trained in agricultural entomology to identify insect vectors of disease. Similarly,
Cambridge-trained, “pure” entomologists drew upon
established British agricultural entomology for their
work in the empire. Insect eradication techniques
constituted the basis of both their medical and agricultural expertise.
When Joseph Chamberlain, as Secretary of State
for the Colonies, sought scientific expertise for his
programme of “constructive imperialism”, Cambridge provided mycologists and entomologists
(Clark, 2001: 99-101). Harold Maxwell Lefroy, who
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achieved the title of “first Imperial Entomologist of
India”, provides perhaps the most instructive example of the relationship between entomological science and empire. He became assistant master of
Seaford College, Sussex after graduating from Cambridge in 1898. Less than a year later, he left this
position to become the entomologist for the newly
created Imperial Department of Agriculture in the
West Indies, where he remained until 1903. He then
succeeded Lionel de Niceville as Entomologist to the
Government of India. Upon the establishment of the
Imperial Agricultural Research Institute at Pusa
(Bihar) in 1905, he became Imperial Entomologist
of India. He subsequently returned to England to
occupy the first chair of entomology at the Imperial
College of Science and Technology, London (Anonymous, 1925b, 1925d; Bateman, 1978). Although
trained in academic entomology at Cambridge,
Lefroy spent his entire career in the service of economic entomology. Aware of his pioneering position
in the conceptual, institutional, and professional
development of his discipline, he brought an evangelical fervour to his subject. His position at Imperial College made him sensitive to his dual audience:
metropolitan and colonial. This, in turn, moulded
him into Britain’s first entomological “researchentrepreneur” (Rosenberg, 1971).
By 1914, economic entomology had achieved status as an acknowledged discipline through the
acquisition of specialist journals, training, and organizations (Lemain et al., 1976; Worboys, 1979: 301;
Whitley, 1982). Its dominance of the nascent Association of Economic Biologists highlighted the subject’s success. Established in 1904 at the instigation
of W.E. Collinge and F.V. Theobald, the Association
effectively drew together all “such Biologists
employed by the Government or by any County or
City Council, University, or Agricultural or Horticultural College or Association...” (Collinge, 1905).
The institutionalization of economic entomology
rested upon emergent agriculturists, Cambridgeeducated imperial zoologists, and high-profile initiatives in tropical medicine. Significantly, the officers
elected at the inaugural meeting of the Association
of Economic Biologists, in November 1904, represented all of these interests. F.V. Theobald became
president. Sir Patrick Manson, A.E. Shipley, a Cambridge zoologist, and William Somerville, a prominent agricultural scientist, shared the vice-presidency. Moreover, economic entomology held a dominant position in the Association. Between 1904 and
1918, all of the presidents were entomologists; and
papers on “entomology and insect pests” constituted the greatest number from any single category
(i.e., 250 of 960) in the first twenty-five volumes of
the Association’s Annals of Applied Biology (19141938) (Brierley, 1939: 178).
Initially institutionalized as an agricultural science, economic entomology sunk its conceptual
roots in the notion of the balance of nature. Posing
as agents of the restoration of a natural equilibrium,
323
economic entomologists used the technological fix
of insecticides to establish their expertise. In a bid
to wrest the lead in medical entomology from medical practitioners, economic entomologists attempted to consolidate the institutional gains that they
had made in agricultural applications. H. Maxwell
Lefroy announced:
Medical men are organised and that so successfully
that in a present problem, largely entomological, the
medical interest has tended to prevent all recognition
of the value and need of the entomologist’s services....
If then the applied biologist is to make himself felt, it
will be through an organisation comparable to those
by which the chemists, the engineers and the doctors
assert themselves; we hope to make the Association
[of Applied Biologists (renamed in 1914)] such an
organised body.... (Lefroy, 1914: 2-3).
Five years later, in 1919, Lefroy pushed for professional closure. He submitted a proposal to convert
the Association from a scientific society to a professional licensing body for applied biologists (Brierley,
1939: 183; Anonymous, 1919-1920). By the early
twentieth century, agricultural science and tropical
medicine, as anti-depressive measures and as “tools
of empire”, had created an increasingly self-conscious body of professional, expert, economic entomologists.
Fly wars
Anti-housefly campaigns of the early twentieth century provided economic entomologists with the
opportunity to address a pressing domestic issue
outside the rubric of tropical medicine. Two principal concerns precipitated unprecedented awareness
of the public health dangers of the housefly: high
infant mortality from summer diarrhoea, and loss of
soldiers’ lives due to typhoid and other enteric disorders. In this respect, the experiences of the Spanish-American and Boer Wars did much to galvanize
the campaign against the housefly. In a commission
headed by army surgeon Walter Reed, an investigation into the outbreak of typhoid fever among
American troops was undertaken in late 1898. Reed
and his team concluded that flies played a significant role in the spread of typhoid by carrying bacteria, contained in faeces, on their wings and feet to
army food. The British Army Medical Department
Report for 1900 drew the same conclusion when it
assessed the outbreak of enteric fever among the
2nd King’s Royal Rifles at Diyatalawa Camp, Ceylon (Austen, 1904).
Similar circumstantial evidence against the fly was
mustered to explain enteric fever at Bloemfontein.
The Boer War focused attention on flies as possible
vectors of typhoid, and it led to the link between
flies and infant diarrhoea. Worried by the poor state
of recruits for the Boer War, a general fear about the
health of next generation gripped the medical community. In particular, they struggled to understand
the rise in infant mortality due to diarrhoeal diseases
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(Dwork, 1987: 1-48). Noting the coincidence in the
seasonal peaks of the disease and housefly populations, medical officers of health – such as J.C.T.
Nash, Arthur Newsholme, James Niven and E.W.
Hope – began to suggest that houseflies spread the
disease from infected excrement to babies’ milk or
mouths. In 1914, it was announced in the House of
Commons that 123 medical officers of health had
warned of the dangers of houseflies in connection
with the summer prevalence of infant diarrhoea. By
the same date, the Local Government Board had
issued five annual reports on the subject, thoroughly investigating the life habits, flight distances and
patterns, and bacterial cultures associated with the
housefly (Graham-Smith, 1913: 252-265). By the
outbreak of the First World War, an anti-fly campaign was in full swing, and the housefly had
become a spreader of filth-germs with legs (Rogers,
1989). Public fear was sufficient for insurance companies to announce policies of assurance covering
the risk of fly-borne infections being carried back
from the battlefields in 1915 (Anonymous, 1915b).
In 1917, the National Baby Week Council received
180,000 essays from schoolchildren for their competition. The rules of the contest perpetuated the
well-established gender assumptions about insect
“hunting”. Boys were instructed to explain “Why I
should kill that fly”, while girls had to recount
“How I mind our baby” (Anonymous, 1917). A war
against the housefly assumed a central place in the
infant welfare movement.
What role did the entomologists play in this campaign? In general, they continued to define themselves as experts in the identification of particular
species of insects, their habits and life histories, and
eradication programmes. In 1907, Robert Newstead, at the behest of E.W. Hope and the Liverpool
Health Authority, produced a special report on the
housefly, replete with graphic photographs of the
eggs, larvae, pupae, and perfect fly in refuse and
assorted dung (Newstead, 1908). The British Museum (Natural History) published Ernest Austen’s The
House-Fly as a Danger to Health as the first in its
series of booklets on applied (economic) entomology in 1913 (Austen, 1913). In addition, a large-scale
model of a housefly, a tray of fly-infested food, and
a heap of kitchen refuse was added to the central
hall display (Anonymous, 1916). And under the
direction of Lefroy and F.M. Howlett, the museum
issued its own “fly danger” poster. The ever-zealous
Lefroy organized an anti-fly exhibition at the Zoological Society’s gardens in 1915, and issued several booklets on the subject (Lefroy, 1915). All were
agreed that the best way of destroying the fly was by
attacking or destroying its breeding places. In this
manner, claimed Shipley, they could repeat the success of anti-malarial, anti-mosquito campaigns – “if
we have the faith which moves mountains - mountains of manure” (Shipley, 1915a: 13). The advent
of war in 1914 saw a large number of horses requisitioned and mobilized (Graham-Smith, 1929: 133).
Both war-time camps and increasingly congested
urban centres were awash with excrement.
Frontline war conditions had long been associated
with the unpleasant company of flies. Writing in
1879, Alfred, Lord Tennyson spelled out the realities of military engagement:
Ever the day with its traitorous death from the loop-holes around
Ever the night with its coffinless corpse to be laid in the ground
Heat like the mouth of a hell, or deluge of cataract skies
Stench of old offal decaying, and infinite torment of flies.
(Tennyson, 1969: 1253)
Trench warfare – with its attendant conditions of
cold and wet, poor diet, and vermin – resulted in
considerable discomfort and sickness. By the second year of the First World War, one author reported that “many officers … fear lice more than they
fear bullets” (Shipley, 1915b: 10). This was the
result of the irritation that the vermin caused, and
the new awareness that they also carried typhus.
But lice were not the only insects to cause nuisance
and disease: flies were considered among the most
significant “minor horrors of war” (Shipley, 1915b:
66-82). A distraught soldier wrote to his mother
from near Ypres: “Soon life will be quite unbearable, and there will be any amount of disease
spread by the flies” (Anonymous, 1915a). The noise
from their buzzing on occasion drowned out the
sound of an approaching shell. One soldier counted
thirty-two dead flies in his shaving water and seventy-two from his shoulder to his wrist. With this
kind of insect company, there were also reports that
snipers would use fly swarms above trenches to
locate potential targets (Winter, 1978: 98-100).
Flies were an undesirable but ubiquitous presence
during war.
Consequently, entomologists, like Newstead,
Lefroy and Austen, took their expertise to sanitary
commissions on the front lines. Members of the
RAMC became very sensitive to the possibility of
the spread of infectious disease with the advent of
war. Initially, they believed that fifteen years of sanitary education had left them well prepared: “no
army was better equipped in knowledge of sanitary
science affecting the field”. They soon feared that
this initial advantage was disappearing with losses
of trained men and their replacement with less
informed new recruits. “Therefore”, it was stated,
“unity in both statement and action of the medical
profession is a factor of all importance...” (Kinner,
1915: 365). Drawing on his experience at Rouen in
1915, Captain P.J. Marett noted that complete lack
of segregation made preventive measures difficult.
Latrines, for instance, were often located next to
kitchens. To address properly the chief “carrier” diseases – enteric and the paratyphoids – a sustained
attack had to be mounted against flies. Breeding
grounds had to be assaulted; flies had to be killed;
and food, latrines, and patients had to be protected
from the unwanted visitations of the winged foes
(Marett, 1915).
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One of the greatest difficulties was the sheer
quantity of excrement. The daily production of
horse dung was estimated at one hundred tons in
Rouen. Under war-time conditions, it was impracticable – if not impossible – to sell the dung as fertiliser. Similarly, it was difficult to burn all of it in
specially constructed incinerators. In the first
instance, heaps were made that were covered with
quicklime, earth, and hay. Some of it was dumped
using a narrow gauge train line that led to a depression, where it was covered with quicklime and
earth, and planted over with flower seeds donated
by Messrs. Carter and Sons. Fly populations were
reduced by treating camps, in general, and latrines,
in particular, with paraffin. Traps of five per cent
formalin in lumps of sugar were suggested; and covers and netting were employed where required. Following the instructions of Lieutenant R. Newstead
of the Entomological Commission, a boiled solution
of five parts castor oil and eight parts resin was
spread on paper or in tins as a substitute for fly
paper. Fly brigades, of one non-commissioned officer and four men, sprayed cookhouses with twoounces-to-the-gallon formalin solutions each week.
In addition, one man vigilantly laid tin traps around
manure heaps and removed clusters of eggs. Marett
estimated that this one man destroyed approximately 236,000 flies, in all their stages of development,
each day (Marett, 1915a).
325
explanation for the increased prevalence of flies was
the recent creation of a dump at Whitlingham
Marshes, half a mile from Postwick’s village church.
They traced the flight of these filthy creatures by
marking them with coloured chalk at the dump, and
trapping them in the village (Copeman et alii,
1911). Through the agency of the fly, rubbish was
revisited upon humanity. The fly traversed boundaries: it transported urban refuse, deposited on rural “wasteland”, back to village homes.
Experts campaigned to change the filthy house-fly’s
name, which, they suggested, was too comfortingly
domestic and innocuous: the public needed to be
alerted to the insidious insects’ danger. Henceforth,
they implored, the housefly should be called the
“typhoid fly” (Howard, 1911: xvi-xvii). Prizes were
offered for the most flies captured and killed in specially designed traps; and major public spectacles
were created to reinforce the link between flies and
infant deaths. In 1915, for instance, Samuel J.
Crumbine, secretary of the Kansas Board of Health
in the USA, organized a sanitation parade, replete
with a huge fly float dragging thirteen black empty
baby buggies (Rogers, 1989: 610; McClary, 1982).
Public health officials were adamant that this campaign was not just about better sanitation. Rather,
they sought to educate and to improve personal
hygiene through active intervention. Health visitors
sought to create more “enlightened mothers”
through educational campaigns and home visits.
Flies and babies
War-time skirmishes with flies and fly-borne diseases intensified campaigns against flies as agents in
the spread of infantile diarrhoea. The deadly Diptera
were frequently shown larger than life on posters, in
films, and in popular magazines. The highly visible
and tangible fly conveniently embodied old notions
of dirt and filth and new fears of unseen germs.
Public health organizations disseminated posters
and songs that denounced flies and filth, and promoted cleanliness. One such song declared:
There was a man in our town
And he was wondrous wise;
He covered up his garbage pail,
To keep away the flies.
(Crew, 1931a)
Similarly, the “Song of the Fly” explained: “The fly
takes a season ticket from the rubbish heap to the
milk jug and other things and this is his song”:
Straight from the rubbish heap I come
I never wash my feet.
And every single chance I get
I walk on what you eat.
(Crew, 1931b)
The Local Government Board undertook a series of
investigations on “Flies as Carriers of Disease” in
the opening decades of the twentieth century. A
“plague of flies” at the village of Postwick, five miles
east of Norwich, provided an opportunity for various experiments. Experts decided that the only
Fear of flies?
Multiple books, solely devoted to the housefly as
disease carrier, were published in the first fifteen
years of the twentieth century. Countless articles on
the same subject appeared in the popular and specialist press. This flurry of interest in the housefly,
however, declined by the end of the second decade
of the twentieth century. In his monograph, The
Housefly (1951), American Luther S. West pondered why almost forty years had elapsed since the
last major study in English of the house-fly. As a historical phenomenon, the campaign to highlight the
housefly peril was part of the infant and child welfare movement, and the rise of a progressive, technocratic push for national efficiency as applied to
public health and to the war effort. For West, this
historical context was not the most important factor
in the scale and extent of fly campaigns at the turn
of the twentieth century: “[O]nce popular interest
had been aroused, imagination tended to outstrip
the facts. The housefly seems to have become
regarded by journalists as the criminal of the ages,
and its elimination or suppression was heralded as
the panacea for most human ills” (West, 1951: ix,
8). Fear, of course, was not unique to the twentieth
century, but the housefly campaign might be
explained within the context of fear borne of the rise
of the “risk society” (Furedi, 1997: 15-44; Beck,
1992). Experts and commentators consistently
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depicted the fly as a product of humanity. Although
it was a creature of nature, the fly had been assured
growth and survival by the rise of urban industrial
society and its attendant problems of overcrowding
and waste.
At a time when the prospect of major mortality
crises, from famine or epidemic disease, seemed to
be a thing of the past in Britain and the USA, the
housefly became the focus for emergent technocrats
who collectively sought to manage lingering threats.
The indeterminate threat of germs carried the same
freight of fear as later mega-hazards, such as genetics and nuclear radiation. Flies were the embodiment of unseen germs: they were “germs with legs”.
The literature of fear that surrounded the fly literally magnified the insect as part of the assessment of
the threat. Illustrations often featured Godzilla-like
flies hovering over cities or over babies. Furthermore, the metaphor of war slipped into the rhetoric
and reality of anti-fly campaigns as the technology
of entomologists and military engineers merged in
the form of insecticides and gases. Identical poisonous gases were used against both insect and human
“enemies” (Russell, 2001; Weindling, 1999).
But fear and bellicose technocrats provide incomplete historical explanations for early twentieth-century anti-housefly campaigns. Similarly, anti-housefly campaigns were not simply emergent malariology transferred to Britain’s temperate shores. Economic entomologists did not acquire knowledge of
insect pests and vectors of disease, and then manufacture the menace of houseflies. Flies, as a significant danger to human health, were a lingering
reminder of a past demographic regime in Britain in
which insect-borne diseases played a significant
role. As James Riley observes, “the insect population
of seventeenth and early eighteenth-century Europe
must have been huge” (Riley, 1986: 852): infrequently washed woolen garments, undrained
swamps, and accumulations of human and animal
waste would have provided ideal feeding and breeding grounds for insect vectors of disease such as
lice, mosquitoes, and flies. The unintended reduction in insect numbers may have played a considerable role in the first phase of post-seventeenth-century European mortality decline. Prior to this
decline, disease arising from filth and insects
accounted for significant mortality among infants,
children, and young adults. Through lavation and
drainage, environmentalist physicians and agricultural improvers reduced the density of insects and
thereby improved the health of people. With rapid
urbanization and urban growth after 1820, however, dense insect populations returned to crowded
urban areas, and the mortality decline stalled until
the early twentieth century. Infant mortality in
Britain, in fact, increased after 1870 when adult
mortality began to decline. The greatest single cause
of infant mortality in towns was diarrhoeal diseases
(Riley, 1986).
Frontline conditions of war concentrated and
magnified sanitary problems with which the British
home-front had struggled for a number of years. A
recent historical case study of a large town, Preston
in Lancashire, contends that an increase in the fly
population was a significant contributing factor in
the anomalous rise in infant mortality in the late
nineteenth century (Morgan, 2002). Increasing
growth of large towns and cities created a new
urban geography that fuelled an increase in horses.
Although the middle-class shift to suburban living
may have been aided by a change in transportation
technology, it generated an expansion of small manufactures and businesses within the city that relied
on horse transportation to service suburban markets. Between 1851 and 1911, the number of horsedrawn carts, vans and wagons in Britain grew from
200,000 to 800,000. Within the commercial sector
alone, the number of horses doubled between 1850
and 1870 and then doubled again between 1870
and 1900: by 1890, Britain’s off-farm horse population exceeded its on-farm population (Morgan,
2002: 102). Each urban horse produced between 15
and 30 pounds of dung and gallons of urine every
day, which provided ideal feeding and breeding
grounds for flies, the vectors of infant diarrhoea. In
Preston in 1880 horses would have produced about
600 tons of manure per square mile. By 1900 this
figure would have risen to 1,600 tons (Morgan,
2002: 116). Under these circumstances, infant mortality in Britain did not begin to decline until the
automobile began to replace the horse and public
health and sanitation reforms became pervasive.
Conclusion
By the early twentieth century, agricultural science,
tropical medicine, and public health had created an
increasingly self-conscious body of professional,
expert, economic entomologists. The house-fly
played a significant role in this process. Unfortunately, for Harold Maxwell Lefroy, the fly also
ensured that he did not live long enough to realize
the full potential of his entomological ambition. In
the end, he succumbed to the thing that he considered his greatest tool in the struggle for professional recognition - insecticides. Eulogized as a “martyr
to the cause of Entomological research” (Anonymous, 1925c: 259), Lefroy, in life and death, epitomized the relationship between insecticides and the
“research-entrepreneur”. He publicly acknowledged
the economic entomologist as a good businessman,
and he made the development of insecticides one of
his chief concerns. In 1916, he announced that his
experiments at Imperial College had produced two
non-poisonous sprays that effectively cleared a
room, hospital ward, kitchen, or any other confined
space of flies. Both of the liquid insecticides were
supplied by the Army Medical Corps. Lefroy made
a clear proprietary claim by naming one of them
“Lefroy’s Solution”; and he declared that he would
not be publishing the formulae “until the require-
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ments of the army are satisfied” (Lefroy, 1916: 10).
Several years earlier, Frank Baines, an architect
from the Office of Works, requested Lefroy’s assistance with an infestation of Death Watch beetle in
the roof of Westminster Hall (Baines, 1914). Lefroy
was one of a number of entomologists to be consulted: C.J. Gahan (British Museum [Natural History]), A.E. Shipley (University of Cambridge), C.
Warburton (Royal Agricultural Society of England),
and Guy Marshall (Imperial Bureau of Entomology)
were also among the group. Collectively, they represented the various strands of the development of
economic entomology in Britain. After experimentation, their proposed solution was an insecticidal liquid. Fortunately, they rejected an initial suggestion
of hydrocyanic gas fumigation in favour of a mixture
of sulphur dioxide and camphor. Most of the developmental work on the appropriate insecticide was
undertaken by two consulting chemists, but the
experience inspired the entrepreneurial spirit in
Lefroy.
In 1924, he and his assistant, Elizabeth Eades,
seized the commercial opportunity that had arisen
and began to supply bottles of woodworm fluid
from a factory in Hatton Garden. Their product –
“Ento-Kill Fluids” – proved so successful that they
attempted to register a company under the same
name the following year. Trade name objections
meant that they had to alter it. Consequently, they
settled for “Rentokil Limited” (Bateman, 1978: 5;
Anonymous, [2006]). Unfortunately, Lefroy’s “flyroom” laboratory at Imperial College of Science and
Technology was improperly ventilated, so he would
never enjoy the fruits of their impending success.
On 10 October 1925, he was overcome by poisonous fumes while experimenting with housefly fumigation techniques with a gas insecticide of his own
invention. He thereby entered “the ranks of those
men who have given their lives in the cause of science...” (Anonymous, 1925a: 899). At an inquest
held later, it was determined that he had rallied
briefly before exposing himself to a final fatal dose.
Aware that his insecticidal research on flies had
proved his undoing, he observed: “The little beggars
got the best of me this time” (Anonymous, 1925e).
On the eve of the Enlightenment, Sir William Temple, diplomat, essayist and mentor of Jonathan Swift,
had contended that flies – wallowing in filth for their
fleeting existence – were the exemplars of modernity
(Hollingsworth, 2001: 134). Post-Enlightenment scientists would, in contrast, associate modernity with
the power of human reason to control and manipulate the natural world. By this accounting, agricultural science, tropical medicine, public health and
sanitation helped to shape an increasingly self-conscious body of modern professional, expert, economic entomologists. Timothy Mitchell, however,
reminds us that despite humanity’s best efforts to
subject the non-human natural world to expertise
and planning, there were always unforeseen or unintended consequences. National politics and scientif-
327
ic expertise, he argues, arose as a bid to manage the
gap between rational human planning and the natural world (Mitchell, 2002: 19-53). By the turn of
the twentieth century, expert economic entomologists had seemingly closed the gap between humanity and nature in their discussions of the housefly
problem: they consistently depicted the fly as a
product of humanity. Although it was a creature of
nature, the fly had been assured growth and survival
by the rise of urban industrial society and its attendant problems of overcrowding and waste. In this
respect, houseflies offer an alternative narrative of
the development of medical entomology in Britain.
Medical entomology, as the study of insects that act
as intermediate hosts for pathogenic organisms,
resulted from developments in parasitology and bacteriology, and from methodological and technical
changes in biomedicine. British involvement in this
process largely occurred in a colonial context, but
environmental conditions in Britain converged to
produce an increased incidence of fly-borne disease
and thereby helped to shape the role of economic
entomologists within the rubric of medical entomology.
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LISTA PARTECI :LISTA PARTECI
26-08-2009
15:48
Pagina 329
List of participants
Baccio Baccetti
Section of Biology
University of Siena
S. Maria alle Scotte Hospital
Lotto 1, Piano 1S
53100 Siena, Italy
baccetti@unisi.it
Mario Coluzzi
Dipartimento di Scienze di Sanità Pubblica
Sezione di Parassitologia
Piazzale Aldo Moro 5
00185 Rome, Italy
mario.coluzzi@uniroma1.it
Jaime L. Benchimol
Casa de Oswaldo Cruz, Fiocruz
Av. Brazil 4365
Manguinhos 21045-900
Rio de Janeiro, RJ, Brazil
jbench@uol.com.br
Jean-Pierre Dedet
CHU Montpellier
Laboratoire de Parasitologie
163 rue Auguste Brossonet
34090 Montpellier, France
parasito@univ-montp1.fr
Karen Brown
Wellcome Unit for the History of Medicine
University of Oxford
45-47 Banbury Road
Oxford OX2 6PE, UK
brown@wuhmo.ox.ac.uk
François Delaporte
Université de Picardie
Faculté de Philosophie et Sciences humaines et
sociales
Chemin du Thil
80025 Amiens Cedex 1, France
delaporte@free.fr
Yves Cambefort
Rehseis, UMR 7596
Université Paris 7
105 rue de Tolbiac
75013 Paris, France
yvecamb@club-internet.fr
Gabriel Gachelin
Rehseis, UMR 7596 CNRS
Université Denis Diderot Paris 7
54 rue de Picpus
75015 Paris, France
ggachel@club-internet.fr
Ernesto Capanna
Dipartimento di Biologia Animale e dell’Uomo
Via A. Borelli 50
00161 Rome, Italy
ernesto.capanna@uniroma1.it
J.F.M. Clark
Institute of Environmental History
University of St Andrews
St Andrews KY16 9AL, Scotland, UK
jfc2@st-andrews.ac.uk
Tamara Giles-Vernick
Unité d’épidémiologie des maladies emergentes
Institut Pasteur
25 rue du Dr Roux
75724 Paris Cedex 15, France
tamara.giles-vernick@pasteur.fr
Dan Gilfoyle
Wellcome Unit for the History of Medicine
University of Oxford
45-47 Banbury Road
Oxford OX2 6PE, UK
daniel.gilfoyle@nationalarchives.gov.uk
LISTA PARTECI :LISTA PARTECI
26-08-2009
330
15:48
Pagina 330
List of participants
Anne Hardy
The Wellcome Trust Centre for the History of
Medicine at UCL
210 Euston Road
London NW1 2BE, UK
ucgaaha@ucl.ac.uk
René Houin
UMR 956 BIPAR
Université Paris XII, Faculté de Médecine
8 rue du général Sarrail
94010 Créteil Cedex, France
houin@noos.fr
Annick Opinel
Centre de recherches historiques
Institut Pasteur
28 rue du Dr Roux
75724 Paris Cedex 15, France
annick.opinel@pasteur.fr
Michael A. Osborne
Department of History
University of California
Santa Barbara, CA 93106-9410, USA
osborne@history.ucsb.edu
Magali Romero Sá
Casa de Oswaldo Cruz, Fiocruz
Av. Brazil 4365
Manguinhos 21045-900 Rio de Janeiro, RJ,
Brazil
magali@fiocruz.br
Michael Worboys
Centre for the History of Science, Technology
and Medicine
Faculty of Life Sciences, University of Manchester
Brunswick Street
Manchester M13 9PL, UK
michael.worboys@manchester.ac.uk

2008 colofon n 4:2005 COLOFON 3/4

Transcription

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