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PALGRAVE STUDIES IN MEDIEVAL AND
EARLY MODERN MEDICINE

Santorio Santori
and the Emergence of
Quantified Medicine,
1614–1790
Corpuscularianism, Technology
and Experimentation

Edited by
Jonathan Barry
Fabrizio Bigotti
 Palgrave Studies in Medieval
and Early Modern Medicine

Series Editors
Jonathan Barry
Department of History
University of Exeter
Exeter, UK

Fabrizio Bigotti
Institute for the History of Medicine
Julius Maximilian University
Würzburg, Germany
The series focuses on the intellectual tradition of western medicine as related to the
philosophies, institutions, practices, and technologies that developed throughout
the medieval and early modern period (500-1800). Partnered with the Centre for
the Study of Medicine and the Body in the Renaissance (CSMBR), it seeks to
explore the range of interactions between various conceptualisations of the body,
including their import for the arts (e.g. literature, painting, music, dance, and
architecture) and the way different medical traditions overlapped and borrowed
from each other. The series particularly welcomes contributions from young
authors. The editors will consider proposals for single monographs, as well as
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(70-120,000 words) or as Palgrave Pivots (upto 50,000 words).

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Alexandra Bamji, University of Leeds
Carmen Caballero-Navas, University of Granada
Klaus-Dietrich Fischer, Johannes Gutenberg-Universität Mainz
David Gentilcore, University of Leicester
Guido Maria Giglioni, University of Macerata
Benjamin Goldberg, University of South Florida
Georgiana Hedesan, University of Oxford
John Henderson, Birkbeck University of London
Martin Kemp, University of Oxford
Ian MacLean, University of Oxford
Cecilia Martini Bonadeo, University of Padua
Heikki Mikkeli, University of Helsinki
William Royall Newman, Indiana University
Vivian Nutton, Centre for the Study of Medicine and the Body in the Renaissance (CSMBR)
Antoine Pietrobelli, Université de Reims Champagne-Ardenne
Aurélien Robert, Centre d’Etudes Supérieures de la Renaissance Tours
Hester Schadee, University of Exeter
Giovanni Silvano, University of Padova
Michael Stolberg, Julius Maximilian University, Würzburg
Alain Touwaide, Institute for the Preservation of Medical Traditions, Washington DC & Los Angeles
Giulia Martina Welston, The Courtauld Institute of Art, London
John Wilkins, University of Exeter
Fabio Zampieri, University of Padova
Fabiola Zurlini, Studio Firmano for the History of Medicine and Science

Editorial Board
Justin Begley, University of Helsinki
Andreas Blank, Alpen-Adria Universität Klagenfurt
Silvana D’Alessio, University of Salerno
Hiro Hirai, Radboud University Nijmegen (Netherlands)
Luca Tonetti, ‘La Sapienza’ University of Rome
Ruben Verwaal, Durham University
Alun Withey, University of Exeter

More information about this series at

http://www.palgrave.com/gp/series/16206
 Jonathan Barry • Fabrizio Bigotti
Editors

Santorio Santori and

the Emergence of
Quantified Medicine,
1614–1790
Corpuscularianism, Technology
and Experimentation
Editors
Jonathan Barry Fabrizio Bigotti
Centre for Medical History Institute for the History of Medicine
University of Exeter Julius Maximilian University
Exeter, UK Würzburg, Germany

ISSN 2524-7387     ISSN 2524-7395 (electronic)

Palgrave Studies in Medieval and Early Modern Medicine
ISBN 978-3-030-79586-3    ISBN 978-3-030-79587-0 (eBook)
https://doi.org/10.1007/978-3-030-79587-0

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 Preface

This book seeks to re-establish the centrality of Santorio Santori,1 not only
in the history of medicine but also in the history of science and technol-
ogy, by showing how, through studying his work and legacy, we obtain a
new and fuller perspective on the nature and development of corpusculari-
anism and early modern experimental philosophy. It does so by establish-
ing not only Santorio’s own contribution to both natural philosophy and
experimentation but also his legacy over the following two centuries, in
which his work was a fundamental reference point to many leading figures.
This legacy, however, was never one of simple acceptance of Santorio’s
ideas and findings: just as he sought to follow Aristotle and Galen by
adopting their methods rather than simply repeating their conclusions, so
successive generations of scholars were inspired to conduct their own pro-
grammes of experimentation and theorising by following his lead, even if
they often sought to explain his results through their own preferred natu-
ral philosophies. Yet, ironically, many of them did so applying versions of
corpuscularian and mechanical philosophy which (though they probably
did not know it) had been pioneered by Santorio himself, although his
own corpuscularian and mechanical models of nature remained largely
implicit in the construction of his instruments and experiments, or were
expressed only briefly in his medical commentaries.
The reader may well wonder why, if Santorio was so important to early
modern science and medicine, has he been so little known or studied in
recent times? One answer might be the paradoxical one that Santorio was
a victim of his own success. His name became indelibly associated with

v
vi Preface

his Medicina statica, which was constantly reprinted, translated and com-
mented on for the next two centuries. In particular his name became syn-
onymous with the weighing chair he invented (the Sanctorian chair) and
with the measurement of insensible perspiration for which it was designed
(widely known as Sanctorian perspiration in the eighteenth century).
Although these remained a living part of scientific and medical theory and
practice until at least the time of Lavoisier, they were sidelined by new
forms of science and medicine in the nineteenth century, and Santorio
became seen as a ‘dead end’ in terms of the progress of science and medi-
cine. Although Lucia Dacome and others have done much to re-establish
the significance of medical statics in the early modern period, they have
tended to present this as an aspect of medical thinking and practice tied
closely to dietetics and the application of the six non-naturals to health
regimes, rather than exploring the broader implications of Santorio’s
work, which this collection seeks to emphasise. The contributions cover
different aspects and developments of Santorio’s legacy throughout
European medicine up to Lavoisier and explore the ‘dissemination’ of his
‘seminal’ ideas. They also demonstrate that Santorio’s researches, both
experimental and theoretical, extended well beyond medicine to cover
theory of matter, optics, clinical practice, technology and even astronomy,
fields in which his contributions served as a fountainhead of new ideas and
pioneered new approaches.
Santorio has also suffered from the downplaying of medicine as a source
of scientific development during the ‘scientific revolution’, and in particu-
lar as a source of mathematical, experimental and ‘mechanical’ models of
nature (including the human body) during this period. Traditionally the
focus has been on physics, and especially the development of a ‘mechanical
worldview’ through development in astronomy associated with men like
Galileo and Newton. More recently, this unilateral view has been rightly
criticised and supplemented by a recognition of the importance of ‘chy-
mistry’ (a term for pre-Lavoiserian chemistry developed by Newman and
Principe in their pioneering work in this area), not least in the work of
Boyle and Newton, which has in turn brought medicine back into the
picture, given the strong links between chymistry and medicine. Historians
have also identified the crucial role played by medical practitioners in the
development of all the sciences, including natural history (e.g. Linnaeus),
while even those natural philosophers not directly practising medicine,
such as Descartes and Leibniz, have been shown to be centrally concerned
with medical developments. Gradually this is filtering into general accounts
of scientific change.
 Preface  vii

As a collection of studies on the development of corpuscularianism and

technology, a clarification as to what we mean by the former term is
required. We have taken corpuscularianism (from the Latin corpusculum
meaning ‘little body’) as that set of theories that explain natural transfor-
mations as the result of the interaction of particles. Most notably, we have
interpreted corpuscularianism as a ‘theory of form’ whereby corpuscles
result from the action of an agent (forma substantialis) that divides the
continuum of matter into portions (corpuscula, particulae) that are pro-
vided with the same specific quality and quantity. Resulting from the divi-
sion of a homogeneous and continuous magnitude, corpuscles can always
be further divided into smaller and smaller parts and for this reason they
are substantially different from atoms and seeds. Thus, as distinct from
both physical and geometrical atomism, corpuscularianism is a develop-
ment of Aristotle’s minima naturalia and it remained a subject of debate
within Aristotelian natural philosophy (as well as among its opponents)
until the late seventeenth century. In this sense, while corpuscularianism is
often associated with the emergence of early modern mechanical philoso-
phy, and especially with the work of Daniel Sennert (1572–1637), René
Descartes (1596–1650) and Robert Boyle (1627–1691), corpuscularian
theories can be found throughout Western philosophy. A new phase of
corpuscularianism occurred in the early modern period (roughly from
Galileo to Newton) when corpuscles were postulated as a necessary aspect
of the mechanical model of the world and thus endowed with properties,
such as shape, discrete quantity and weight. This represented a transition
from the Aristotelian physics of qualities towards a rigorous atomism. But
during this period many authors conflate together different aspects of the
two traditions in ways that are peculiar to their approach to the continuum
and the elemental composition of matter.
Although coming to full bloom in the seventeenth century, the trend
towards the mathematisation of forms started in the late fourteenth cen-
tury. In order to be responsive to mathematical treatment, matter ought
to be particulate and divisible into minima of time, intensity, space, light,
motion and so on. This ‘mathematical minimism’ opened up the possibility
of reinterpreting the metaphysical concept of form, which was redefined as
a structure or ‘geometrical configuration’ whereby forms are reduced to
numerical entities and the emergence of new properties is seen as the
result of a different spatial arrangement of the material substratum. Early
attempts to develop this new idea, however, were tied up to the Aristotelian
conception of place, which requires the existence of absolute directions,
viii Preface

and thus retained the concept of form as an active principle able to guide
the motion of corpuscles in an orderly manner. Such attempts are part of
what has been defined as an ‘Aristotelian corpuscularianism’ predomi-
nantly linked to North Italian philosophers (especially in Padua), includ-
ing Santorio. Compared to other Aristotelians, these thinkers, mainly
physicians, upheld a more direct commitment to the physical existence of
minima in their study of the combination of artificial substances to pursue
a truly quantitative and chymical analysis of them. In many cases, however,
they remained committed to the existence of substantial forms which they
used to explain the emergence of new properties. Santorio’s own version
of corpuscularianism and the adaptations and responses to this found in
the other natural philosophers reveal the complexity of this process.
This volume explores one particular aspect of this new approach by
detailing Santorio’s approach to medicine in the light of the theories he
developed, the instruments he invented and the experimental practices
he pioneered. It collects papers resulting from the international confer-
ence on ‘Humours, Mixtures and Corpuscles. A Medical Approach to
Corpuscularianism in the Seventeenth Century’ that we organised in
Pisa in May 2017 with support from the Wellcome Trust (Grant no.
WT106580/Z/14/Z) and the Institutio Santoriana—Fondazione Comel.
Bigotti’s contributions on Santorio himself, and the chapters on Obizzi, an
early opponent (by Zurlini), and on Santorio’s views on plague (D’Alessio
and Nutton), reveal both his continued commitment to a model of nature
which underpinned a medicine delivered with certainty by the rational phy-
sician and the fundamental departures from traditional medical orthodoxy
which his approach produced. The subsequent chapters of this volume
explore these themes for such key figures as Sennert (Newman), Beeckman
(Moreau), Descartes (Baldassarri), Leibniz (Blank) and Boyle (Ricciardo).
It is clear that Santorio himself was read differently depending on the par-
ticular approach of each author and indeed our contributors (like their
subjects) take rather different positions on the degree to which a corpuscu-
larian approach presumed, for example, the rejection of substantial forms.
With the establishment by the end of the century of forms of ‘mechani-
cal philosophy’ such as Newtonianism and the associated iatromechanical
model of the body/medicine, Santorio ceased, for a period, to be a direct
source of inspiration for theories of matter as a whole. However, his math-
ematical approach to the study of the body and his version of the notion
of ‘insensible perspiration’ remained very important to experimental tra-
ditions within medicine and natural philosophy, as is shown by the later
chapters on Borelli (Zampieri), Baglivi (Tonetti), De Gorter (Verwaal) and
 Preface  ix

Linnaeus (Thomaz). The work of Lavoisier and Séguin (explored in the

final chapter by Antonelli) also shows how Santorio’s legacy could take on
new life and meaning as a new form of chemistry was forged.
In addition to its substantive contributions, we also hope that this vol-
ume will contribute methodologically, both to the historiography that
places medicine at the centre of broader scientific developments in the
early modern period and to those approaches which stress the complexity
of how both old and new models and practices were combined, recovering
the significance of figures such as Santorio who may not fit neatly into
paradigms of ‘scientific revolution’ as marked by dramatic changes of
worldview, but nevertheless reveal that more incremental changes can nev-
ertheless embody significant new approaches that underpin crucial fea-
tures of our own understandings of nature, the body, and medicine.

Exeter, UK Jonathan Barry

Würzburg, Germany  Fabrizio Bigotti

Note

1. We have chosen Santori as the most accurate rendering of his surname, as

found in original documents letters, but he is often called Santorio Santorio
or simply Sanctorius. However, we have used Santorio as the short form of
his name in line with how Galileo’s name is usually rendered in English
forms of his titles (Galileo Galilei = Galileo).
 Acknowledgments

The introduction (by Bigotti and Barry) was made possible by research
funded by the Wellcome Trust under grants 106580/Z/14/Z—Wellcome
Research Fellowship for Bigotti entitled ‘Santorio Santori and the
Emergence of Quantifying Procedures in Medicine at the end of the
Renaissance: Problems, Context, Ideas’—and grant SIA
097782/Z/11/Z—Wellcome Senior Investigator in the Medical
Humanities award for Barry for his project ‘The Medical World of Early
Modern England, Wales and Ireland, c1500–1715’. The chapter by
Bigotti was made possible by research funded by the Wellcome Trust
under grants 106580/Z/14/Z—Wellcome Research Fellowship for
Bigotti entitled ‘Santorio Santori and the Emergence of Quantifying
Procedures in Medicine at the end of the Renaissance: Problems,
Context, Ideas’.

xi
 Contents

1 Introduction  1
Fabrizio Bigotti and Jonathan Barry
1 A Tale of Oblivion and Rebirth  2
2 Santorio’s Life and Works  5
3 ‘Not that Close’: The Problematic Relations Between
Santorio and Galileo 26
4 New Instruments for a New Medicine 34
5 Outlines for a Conclusion 38

2 ‘Gears of an Inner Clock’: Santorio’s Theory of Matter

and Its Applications 65
Fabrizio Bigotti
1 Purpose, Context and Development of Santorio’s Natural
Philosophy 66
2 The Architecture of the Theory 68
3 Limits, Strengths and Applications 80
4 Conclusions 90

3 The Uncertainty of Medicine: Readings and Reactions

to Santorio Between Tradition and Reformation
(1615–1721)103
Fabiola Zurlini
1 The Philosophical and Cultural Backdrop of Obizzi’s
Polemic105

xiii
xiv Contents

2 Obizzi’s Motifs and Arguments in the ‘Staticomastix’106

3 Leonardo Di Capua: Uncertainty as Intrinsic to Medical
Practice110
4 Santorio in England: Popular and Learned Criticisms111
5 Conclusions113

4 Daniel Sennert’s Response to Santorio Santori in the

Light of Chymical Atomism119
William R. Newman
1 Atomism and Occult Qualities119
2 Sennert Versus Santorio126
3 Conclusion130

5 Atoms, Mixture, and Temperament in Early Modern

Medicine: The Alchemical and Mechanical Views of
Sennert and Beeckman137
Elisabeth Moreau
1 Sennert on Minimal Particles and the Superior Form140
2 Isaac Beeckman on Atomic Elements and Geometrical
Proportion147
3 Santorio’s Theory of Mixture in Light of Sennert and
Beeckman153
4 Conclusion156

6 Santorio, Regius, and Descartes: The Quantification and

Mechanization of the Passions in Seventeenth-Century
Medicine165
Fabrizio Baldassarri
1 Henricus Regius Between Santorio and Descartes167
2 Regius Against Descartes: The Status of the Mind171
3 Descartes’ Mechanics of Passions172
4 Regius on Passions174
5 Santorio: Weighing the Passions176
6 Conclusion: A Complementary Association179
 Contents  xv

7 Santorio and Leibniz on Natural Immortality: The

Question of Emergence and the Question of Emanative
Causation191
Andreas Blank
1 Introduction191
2 Natural Immortality and the Question of Emergence194
3 Natural Immortality and the Question of Emanative
Causation203
4 Conclusion208

8 Santorio Santori on Plague: Ideas and Experience

Between Venice and Naples217
Vivian Nutton and Silvana D’Alessio

9 “An inquisitive man, considering when and where he

liv’d”: Robert Boyle on Santorio Santori and Insensible
Perspiration239
Salvatore Ricciardo
1 Introduction239
2 Boyle’s Early Atomism and Santorio245
3 The Doctrine of Effluvia and Boyle’s Corpuscular
Philosophy247
4 The Human Body and Insensible Perspiration: Between
Chymistry and Mechanics250
5 Experimenting on Insensible Perspiration256
6 Epilogue257

10 Giovanni Alfonso Borelli and Santorio on the

Explanation of Fevers273
Fabio Zampieri
1 Introduction: Borelli’s Life and Work273
2 Borelli’s Work on Pestilential Fevers277
3 Conclusion284
xvi Contents

11 Bodies in Balance: Santorio’s Legacy in Baglivi’s Medicine289

Luca Tonetti
1 Introduction289
2 A New Interpretation of Santorio’s Statics: Towards a
Fibrillary Conception of Human Body294
3 Does a “statica mentis” Exist?301
4 Conclusions305

12 Disputing Santorio: Johannes de Gorter’s Neurological

Theory of Insensible Perspiration317
Ruben E. Verwaal
1 Balancing Ingestion and Excretion320
2 Perspiration and the Nerves323
3 Spirits and Effluvia329
4 Sweating it Out334

13 Santorio’s Influence on the Dietetics of Carl Linnaeus347

Luciana Costa Lima Thomaz
1 A Work to Be Kissed347
2 On the Development of Carl Linnaeus’ Medical
Thinking348
3 Santorio in Linnaeus’ Dietetic353
4 Linked by Aphorisms363
5 Conclusions364

14 Weighing Authority: Lavoisier’s and Séguin’s

Reassessment of Santorio’s Experiments
on Transpiration371
Francesca Antonelli
1 Lavoisier’s Chemical Education and Medicine371
2 Lavoisier and Santorio376
3 The “chemical” Physiology of Respiration and
Transpiration: Lavoisier’s and Séguin’s Response to the
Sanctorian Tradition382
4 Conclusion390

Index403
 Notes on Contributors

Francesca Antonelli is currently research fellow at the University of

Bologna. In May 2021 she obtained a PhD in History of Science at the
University of Bologna and the École des Hautes Études en Sciences
Sociales, with a thesis on Marie-Anne Paulze-Lavoisier (1758–1836) and
Lavoisier’s Registres de laboratoire. Her main research interests concern
scientific practices, sociability, and gender in the long eighteenth century.
Fabrizio Baldassarri is a Marie Skłodovska Curie fellow at Ca’ Foscari
University of Venice and Indiana University Bloomington. His research
focuses on early modern natural philosophy, with an emphasis on the nat-
uralistic studies of Descartes, plants, and the early modern life sciences. He
has been a post-doctoral researcher at the University of Bucharest, at
Gotha Centre, at Bar-Ilan University in Tel Aviv, at Utrecht
University, at HAB in Wolfenbüttel, and has published on early mod-
ern natural philosophy, botany, medicine, and sciences
Jonathan Barry is an Emeritus Professor of History at the University of
Exeter, UK, and guest professor at Ludwig Maximilian University (LMU)
in Munich, Germany. He is an early modern social and cultural historian
with a particular interest in the history of science, medicine, and witchcraft
and co-editor of Palgrave Historical Studies in Witchcraft and Magic. He
was co-founder of the Centre for Medical History at Exeter and a Wellcome
Senior Investigator in Medical Humanities (2012–2018) for a project on
medical practice in England, Ireland, and Wales 1500–1715.

xvii
xviii Notes on Contributors

Fabrizio Bigotti is the Director of the Centre for the Study of Medicine
and the Body in the Renaissance (CSMBR), as well as a research fellow at
the Institute for the History of Medicine at the Julius Maximilian
University of Würzburg in Germany and an honorary Research Fellow at
the College of Humanities at the University of Exeter, UK. As an intel-
lectual historian, he specialises in the history of science, medicine, and
technology of the late medieval and early modern period (1300–1700),
focusing particularly on on the history of quantification and the role that
classical and medieval philosophies played in the development of early
modern logic, theory of matter, anatomy, and physiology. His publications
include works on late Renaissance Galenism, Vesalius, Acquapendente,
Santorio Santori, Scholasticism, early modern corpuscularianism and the
invention of precision instruments.
Andreas Blank holds a research position funded by the Austrian Science
Fund (FWF) at Alpen-Adria Universität Klagenfurt. He has been a visiting
associate professor at the University of Hamburg and Bard College Berlin
and visiting fellow at the Center for Philosophy of Science (University
of Pittsburgh), the Cohn Institute for the History and Philosophy of
Science and Ideas (Tel Aviv University), and the Istituto per il Lessico
Intellettuale e Storia delle Idee (CNR, Rome). He is the author of
some 80 articles, mainly on early modern philosophy, in edited vol-
umes and journals such as Annals of Science, British Journal for the
History of Philosophy, Early Science and Medicine, Eighteenth-Century
Studies, European Journal of Philosophy, History of European Ideas, History
of Philosophy Quarterly, Hypatia, Intellectual History Review, Journal of
Early Modern Studies, Journal of Modern Philosophy, Journal of the History
of Ideas, Journal of International Political Theory, The Monist, Perspectives
on Science, Science in Context, and Studia Leibnitiana.
Silvana D’Alessio is Professor of Modern History at the University of
Salerno; her main interests are the revolt in Naples in 1647–1648, the
plague in the kingdom of Naples in 1656 and Hippocratic medicine in
the early modern age (especially in Pietro Andrea Canoniero’s treatises).
Her main works on medical themes are Per un principe “medico publico”.
Il percorso di Pietro Andrea Canoniero (Scandicci: CET, 2015); “Usi
politici della medicina nella prima età moderna”, in Interpretare e curare.
Medicina e salute nel Rinascimento, ed. by M. Conforti, A. Carlino and
A. Clericuzio, Rome, Carocci, 2013, pp. 269–282; “On the Plague in
Naples, 1656: Expedients and Remedies” in C. De Caprio, D. Cecere,
 Notes on Contributors  xix

L. Gianfrancesco, P. Palmieri (eds.), Disaster Narratives in Early Modern

Naples. Politics, Communication and Culture, 2018, pp. 187–204; “L’aria
innocente. Geronimo Gatta e le sue fonti,” in Mediterranea-ricerche
storiche, a. XV, n. 44, dic. 2018, pp. 587–612; “Un allievo di Marco
Aurelio Severino sulla peste di Napoli,” Medicina Historica, vol. 4 (1).
Elisabeth Moreau is an FNRS Postdoctoral Researcher at the Université
Libre de Bruxelles in Belgium. Trained in history and philosophy of sci-
ence, she started her postdoctoral research at Princeton University in
2019. Her project, “From the Alembic to the Stomach: Nutrition and
Pharmacology in Early Modern Medicine,” is centred on the medical and
alchemical conceptions of digestion between 1550 and 1650 in Europe.
Previous to this project, her doctoral dissertation focused on the emer-
gence of atomistic and corpuscular theories in late Renaissance Galenic
medicine.
William R. Newman is Distinguished Professor and Ruth N. Halls
Professor in the Department of History and Philosophy of Science
at Indiana University. Most of his work in the history of science has been
devoted to alchemy and “chymistry,” the art-nature debate, and matter
theories, particularly atomism. Newman is also General Editor of
the Chymistry of Isaac Newton, an online resource combining born-­
digital editions of Newton’s alchemical writings with multimedia replica-
tions of Newton’s alchemical experiments. In addition, he was Director of
the Catapult Center for Digital Humanities and Computational Analysis
of Texts at Indiana University. He is on the editorial boards
of Archimedes, Early Science and Medicine, and HOPOS.
Vivian Nutton is an Emeritus Professor of the History of Medicine at
University College London. A Cambridge classicist by training, he devel-
oped his interest in the Renaissance when he moved to the then Wellcome
Institute in 1977. Since then he has written widely on almost all aspects of
medicine from Antiquity to the seventeenth century. His most recent pub-
lications include annotated translations of Johann Guinter and Andreas
Vesalius’ Principles of Anatomy (2017) and of John Caius’ autobibliogra-
phy (2018), as well as a study of Galen of Pergamum (2020). His
Renaissance medicine. A short history of medicine in the sixteenth century is
scheduled for publication in 2022. As well as being a member of learned
Academies around Europe, he is also a member of the Ancient Society of
College Youths of London.
xx Notes on Contributors

Salvatore Ricciardo teaches history of science at the University of

Bergamo, Department of Human and Social Sciences. He has published
articles and books on early modern science and philosophy, and he is
­co-­author of “The Reappearance of Galileo’s Original Letter to Benedetto
Castelli” in Notes and Records. The Royal Society Journal of the History of
Science, 2018. His research interests include the relations between science
and religion, the seventeenth-century debates over the theory of matter,
and the seventeenth-century Italian scientific academies.
Luciana Costa Lima Thomaz is a physician, specialist in the field of
integrative medicine. Her master’s and doctoral research, developed in the
History of Science programme at the Pontifical University of São Paulo,
focused on the history of medicine, especially in complementary and alter-
native medicine in the twentieth century. Her postdoctoral research,
developed in the same programme in association with the Department of
History of Science and Ideas at the University of Uppsala, concerned the
medicine of Carl von Linné and his particular way of observing the physi-
ology of the human body.
Luca Tonetti is currently Research Fellow in History of Science at
University of Bologna. He received his PhD in History and Philosophy of
Science from Sapienza University of Rome, with a dissertation on Giorgio
Baglivi and his reform of medical practice in De praxi medica (1696). He
has since held a postdoctoral fellowship at the Centre d’Études Supérieures
de la Renaissance (CESR), University of Tours, and at the Herzog August
Bibliothek, Wolfenbüttel. Since 2017 he is a member of the editorial team
of Nuncius. Journal of the Material and Visual History of Science (Brill),
and is now serving as Book Review Editor. His work focuses on the his-
tory of medicine, particularly on anatomy and medical practice in Italy in
the seventeenth and eighteenth centuries.
Ruben E. Verwaal (PhD, 2018) is a Dutch Research Council Rubicon
Research Fellow at the Institute for Medical Humanities, Durham
University, and curator of the medical collections at Erasmus University
Medical Centre, Rotterdam. He has a particular interest in the history and
material culture of early modern science and medicine. His first book
is titled Bodily Fluids, Chemistry and Medicine in the Eighteenth-Century
Boerhaave School (Palgrave Macmillan, 2020). Verwaal is working on
the medical perceptions and personal experiences of deafness and hardness
of hearing in early modern Europe.
 Notes on Contributors  xxi

Fabiola Zurlini is Vice-Director and Director in Chief of the Studio

Firmano for the History of Medicine and Science (Fermo). Her research
interests and publications focus on the early modern history of medical
libraries and medical bibliography, medical education, and the medical
profession, with a special focus on medicine at the Roman court of the
Queen Christina of Sweden. She is working on publications devoted to
the physicians working at the Roman Court of Queen Christina.
Another random document with
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 CHAPTER XVII.

THE DISTRIBUTION OF FRESHWATER FISHES.

Having shown above that numerous marine fishes enter fresh

waters, and that some of them have permanently established
themselves therein, we have to eliminate from the category of
freshwater fishes all such adventitious elements. They are derived
from forms, the distribution of which is regulated by other agencies,
and which, therefore, would obscure the relations of the faunæ of
terrestrial regions if they were included in them. They will be
mentioned with greater propriety along with the fishes constituting
the fauna of the brackish water.
True freshwater fishes are the following families and groups only:
—
Dipnoi with 4 species.
Acipenseridæ and „ 26 „
Polyodontidæ
Amiidæ „ 1 „
Polypteridæ. „ 2 „
Lepidosteidæ. „ 3 „
Percina „ 46 „
Grystina „ 11 „
Aphredoderidæ „ 1 „
Centrarchina „ 26 „
Dules „ 10 „
Nandidæ „ 7 „
Polycentridæ „ 3 „
Labyrinthici „ 30 „
Luciocephalidæ „ 1 „
Gastrosteus „ 10 „
Ophiocephalidæ „ 31 „
Mastacembelidaæ „ 13 „
Chromides „ 105 „
Comephoridæ „ 1 „
Gadopsidæ „ 1 „
Siluridæ „ 572 „
Characinidæ „ 261 „
Haplochitonidæ „ 3 „
Salmonidæ (3 genera excepted) „ 135 „
 Percopsidæ „ 1 „
Galaxiidæ „ 15 „
Mormyridæ (and Gymnarchidæ) „ 52 „
Esocidæ „ 8 „
Umbridæ „ 2 „
Cyprinodontidæ „ 112 „
Heteropygii „ 2 „
Cyprinidæ „ 724 „
Kneriidæ „ 2 „
Hyodontidæ „ 1 „
Osteoglossidæ „ 5 „
Notopteridæ „ 5 „
Gymnotidæ „ 20 „
Symbranchidæ „ 5 „
Petromyzontidæ „ 12 „
Total 2269 species.

As in every other class of animals, these freshwater genera and

families vary greatly with regard to the extent of their geographical
range; some extend over the greater half of the continental areas,
whilst others are limited to one continent only, or even to a very small
portion of it. As a general rule, a genus or family of freshwater fishes
is regularly dispersed and most developed within a certain district,
the species and individuals becoming scarcer towards the periphery
as the type recedes more from its central home, some outposts
being frequently pushed far beyond the outskirts of the area
occupied by it. But there are not wanting those remarkable instances
of closely allied forms occurring, almost isolated, at most distant
points, without being connected by allied species in the intervening
space; or of members of the same family, genus, or species
inhabiting the opposite shores of an ocean, and separated by many
degrees of abyssal depths. We mention of a multitude of such
instances the following only:—

A. Species identical in distant continents—

1. A number of species inhabiting Europe and the temperate
parts of eastern North America, as Perca fluviatilis, Gastrosteus
pungitius, Lota vulgaris, Salmo solar, Esox lucius, Acipenser sturio,
Acipenser maculosus, and several Petromyzonts.
 2. Lates calcarifer is common in India as well as in Queensland.
3. Galaxias attenuatus inhabits Tasmania, New Zealand, the
Falkland Islands, and the southernmost part of the South American
continent.
4. Several Petromyzonts enter the fresh waters of Tasmania,
South Australia, New Zealand, and Chili.

B. Genera identical in distant continents—

1. The genus Umbra, so peculiar a form as to be the type of a
distinct family consisting of two most closely allied species only, one
of which is found in the Atlantic States of North America, the other in
the system of the Danube.
2. A very distinct genus of Sturgeons, Scaphirhynchus, consisting
of two species only, one inhabiting fresh waters of Central Asia, the
other the system of the Mississippi.
3. A second most peculiar genus of Sturgeons, Polyodon,
consists likewise of two species only, one inhabiting the Mississippi,
the other the Yang-tse-kiang.
4. Amiurus, a Siluroid, and Catostomus, a Cyprinoid genus, both
well represented in North America, occur in a single species in
temperate China.
5. Lepidosiren is represented by one species in tropical America,
and by the second in tropical Africa (Protopterus).
6. Notopterus consists of three Indian and two West African
species.
7. Mastacembelus and Ophiocephalus, genera characteristic of
the Indian region, emerge severally by a single species in West and
Central Africa.
8. Symbranchus has two Indian and one South American
species.
9. Prototroctes, the singular antarctic analogue of Coregonus,
consists of two species, one in the south of Australia the other in
New Zealand.
10. Galaxias is equally represented in Southern Australia, New
Zealand, and the southern parts of South America.

C. Families identical in distant continents—

1. The Labyrinthici, represented in Africa by 5, and in India by 25
species.
2. The Chromides, represented in Africa by 25, and in South
America by 80 species.
3. The Characinidæ, represented in Africa by 35, and in South
America by 226 species.
4. The Haplochitonidæ, represented in Southern Australia by
one, in New Zealand by one, and in Patagonia by a third species.
This list could be much increased from the families of Siluridæ
and Cyprinidæ, but as these have a greater range than the other
Freshwater fishes, they do not illustrate with equal force the object
for which the list has been composed.

The ways in which the dispersal of Freshwater fishes has been

effected were various; they are probably all still in operation, but
most work so slowly and imperceptibly as to escape direct
observation; perhaps, they will be more conspicuous, after science
and scientific inquiry shall have reached to a somewhat greater age.
From the great number of freshwater forms which we see at this
present day acclimatised in, gradually acclimatising themselves in, or
periodically or sporadically migrating into, the sea, we must conclude
that, under certain circumstances, salt water may cease to be an
impassable barrier at some period of the existence of freshwater
species, and that many of them have passed from one river through
salt water into another. Secondly, the headwaters of some of the
grandest rivers, the mouths of which are at opposite ends of the
continents which they drain, are sometimes distant from each other a
few miles only; the intervening space may have been easily bridged
over for the passage of fishes by a slight geological change affecting
the level of the watershed, or even by temporary floods; and a
communication of this kind, if existing for a limited period only, would
afford the ready means of an exchange of a number of species
previously peculiar to one or the other of those river or lake systems.
Some fishes, provided with gill-openings so narrow that the water
moistening the gills cannot readily evaporate; and endowed,
besides, with an extraordinary degree of vitality, like many Siluroids
(Clarias, Callichthys), Eels, etc., are enabled to wander for some
distance over land, and may thus reach a water-course leading them
thousands of miles from their original home. Finally, fishes or their
ova may be accidentally carried by waterspouts, by aquatic birds or
insects, to considerable distances.
Freshwater fishes of the present fauna were already in existence
when the great changes of the distribution of land and water took
place in the tertiary epoch; and having stated that salt water is not an
absolute barrier to the spreading of Freshwater fishes, we can now
more easily account for those instances of singular disconnection of
certain families or genera. It is not necessary to assume that there
was a continuity of land stretching from the present coast of Africa to
South America, or from South America to New Zealand and
Australia, to explain the presence of identical forms at so distant
localities; it suffices to assume that the distances were lessened by
intervening archipelagoes, or that an oscillation has taken place in
the level of the land area.
Dispersal of a type over several distant continental areas may be
evidence of its great antiquity, but it does not prove that it is of
greater antiquity than another limited to one region only. Geological
evidence is the only proof of the antiquity of a type. Thus, although
the Dipnoi occur on the continents of Africa, South America, and
Australia, and their present distribution is evidently the consequence
of their wide range in palæozoic and secondary epochs; the proof of
their high antiquity can be found in their fossil remains only. For,
though the Siluroids have a still greater range, their wide distribution
is of comparatively recent date, as the few fossil remains that have
been found belong to the tertiary epoch. The rapidity of dispersal of a
type depends entirely on its facility to accommodate itself to a variety
of physical conditions, and on the degree of vitality by which it is
enabled to survive more or less sudden changes under unfavourable
conditions; proof of this is afforded by the family of Siluroids, many of
which can suspend for some time the energy of their respiratory
functions, and readily survive a change of water.

To trace the geological sequence of the distribution of an ichthyic

type, and to recognise the various laws which have governed, and
are still governing its dispersal, is one of the ultimate tasks of
Ichthyology. But the endeavour to establish by means of our present
fragmentary geological knowledge the divisions of the fauna of the
globe, leads us into a maze of conflicting evidence; or, as Mr.
Wallace truly observes, “any attempt to exhibit the regions of former
geological ages in combination with those of our own period must
lead to confusion.” Nevertheless, as the different types of animals
found at the present day within a particular area have made their
appearance therein at distant periods, we should endeavour to
decide as far as we can, in an account of the several zoo-
geographical divisions, the following questions:—
1. Which of the fishes of an area should be considered to be the
remnants of ancient types, probably spread over much larger areas
in preceding epochs?
2. Which of them are to be considered to be autochthont species,
that is, forms which came in the tertiary epoch or later into existence
within the area to which they are still limited, or from which they have
since spread?
3. Which are the forms which must be considered to be
immigrants from some other region?
The mode of division of the earth’s surface into zoological regions
or areas now generally adopted, is that proposed by Mr. Sclater,
which recommends itself as most nearly agreeing with the
geographical divisions. These regions are as follows:—
I. Palæogæa.
1. The Palæarctic region; including Europe, temperate Asia,
and North Africa.
2. The Ethiopian region; including Africa, south of the
Sahara, Madagascar, and the Mascarene Islands; also
Southern Arabia.
3. The Indian region; including India south of the
Himalayas, to Southern China, Borneo, and Java.
4. The Australian region; including Australia, the Pacific
Islands, Celebes, and Lombock.
II. Neogæa.
5. The Nearctic region; including North America to Northern
Mexico.
6. The Neotropical region; including South America, the
West Indies, and Southern Mexico.
Comparatively few classes and orders of animals have been
carefully studied with regard to their geographical distribution, but the
majority of those which have been examined show that the
difference of latitude is accompanied by a greater dissimilarity of
indigenous species than that of longitude, and that a main division
into an old world and new world fauna is untenable. More especially
the Freshwater fishes, with which we are here solely concerned,
have been spread in circumpolar zones, and in a but limited degree
from north to south. No family, much less a genus, ranges from the
north to the south, whilst a number of families and genera make the
entire circuit, and some species more than half of the circuit round
the globe within the zone to which they belong. Not even the
Cyprinoids and Siluroids, which are most characteristic of the
freshwater fauna of our period, are an exception to this. Temperature
and climate, indeed, are the principal factors by which the character
of the freshwater fauna is determined; they form the barriers which
interfere with the unlimited dispersal of an ichthyic type, much more
than mountain ranges, deserts, or oceans. Hence the tropical zone is
an impassable barrier to the northern Freshwater fish in its progress
towards the south; where a similarly temperate climate obtains in the
southern hemisphere, fish-forms appear analogous to those of the
north, but genetically and structurally distinct.
The similarity which obtains in fishes at somewhat distant points
of the same degree of longitude, rarely extends far, and is due to the
natural tendency of every animal to spread as far as physical
conditions will permit. Between two regions situated north and south
of each other there is always a debateable border ground, in parts of
which sometimes the fishes of the one, sometimes those of the
other, predominate, and which is, in fact, a band of demarcation.
Within this band the regions overlap each other; therefore, their
border lines are rarely identical, and should be determined by the
northern and southernmost extent of the most characteristic types of
each region. Thus, for instance, in China, a broad band intervenes
between temperate and tropical Asia, in which these two faunæ mix,
and the actual northern border line of the tropical fauna is north of
the southern border line of temperate Asia.
It is the aim of every philosophical classification to indicate the
degree of affinity which obtains between the various divisions; but
the mode of division into six equivalent regions, as given above,
does not fulfil this aim with regard to Freshwater fishes, the
distribution of which allows of further generalisation and subdivision.
The two families, Cyprinidæ and Siluridæ, of which the former yields
a contingent of one-third, and the latter of one-fourth of all the
freshwater species known of our period, afford most valuable
guidance for the valuation of the degrees of affinity between the
various divisions. The Cyprinoids may be assumed to have taken
their origin in the Alpine region, dividing the temperate and tropical
parts of Asia; endowed with a greater capability of acclimatising
themselves in a temperate as well as tropical climate than any other
family of freshwater fishes, they spread north and south as well as
east and west; in the preglacial epoch they reached North America,
but they have not had time to penetrate into South America,
Australia, or the islands of the Pacific. The Siluroids, principally
fishes of the sluggish waters of the plains, and well adapted for
surviving changes of the water in which they live, for living in mud or
sea-water, flourish most in the tropical climate, in which this type
evidently had its origin. They came into existence after the
Cyprinoids, fossil remains being known only from tertiary deposits in
India, none from Europe. They rapidly spread over the areas of land
within the tropical zone, reaching northern Australia from India, and
one species even immigrated into the Sandwich Islands, probably
from South America. The Coral Islands of the Pacific still remain
untenanted by them. Their progress into temperate regions was
evidently slow, only very few species penetrating into the temperate
parts of Asia and Europe; and the North American species, although
more numerous, showing no great variety of structure, all belonging
to the same group (Amiurina). Towards the south their progress was
still slower, Tasmania, New Zealand, and Patagonia being without
representatives, whilst the streams of the Andes of Chili are
inhabited by a few dwarfed forms identical with such as are
characteristic of similar localities in the more northern and warmer
parts of the South American continent.
After these preliminary remarks we propose the following division
of the fauna of Freshwater fishes:—
I. The Northern Zone.—Characterised by Acipenseridæ. Few
Siluridæ. Numerous Cyprinidæ. Salmonidæ, Esocidæ.
1. Europo-Asiatic or Palæarctic Region.—Characterised by
absence of osseous Ganoidei; Cobitidæ and Barbus
numerous.
2. North American Region.—Characterised by osseous Ganoidei,
Amiurina, and Catostomina; but no Cobitidæ or Barbus.
II. The Equatorial Zone.—Characterised by the development of
Siluridæ.
A. Cyprinoid Division.—Characterised by presence of Cyprinidæ
and Labyrinthici.
1. Indian Region.—Characterised by [absence of Dipnoi[18]]
Ophiocephalidæ, Mastacembelidæ. Cobitidæ numerous.
 2. African Region.—Characterised by presence of Dipnoi and
Polypteridæ. Chromides and Characinidæ numerous.
Mormyridæ. Cobitidæ absent.
B. Acyprinoid Division.—Characterised by absence of Cyprinidæ
and Labyrinthici.
1. Tropical American Region.—Characterised by presence of
Dipnoi. Chromides and Characinidæ numerous.
Gymnotidæ.
2. Tropical Pacific Region.—Characterised by presence of
Dipnoi. Chromides and Characinidæ absent.
III. The Southern Zone.—Characterised by absence of Cyprinidæ,
and scarcity of Siluridæ. Haplochitonidæ and Galaxiidæ
represent the Salmonoids and Esoces of the Northern zone.
One region only.
1. Antarctic Region.—Characterised by the small number of
species; the fishes of—
a. The Tasmanian sub-region; b. The New Zealand sub-region;
c. The Patagonian sub-region;
being almost identical.[19]
In the following detailed account we begin with a description of
the equatorial zone, this being the one from which the two principal
families of freshwater fishes seem to have spread.

I. Equatorial Zone.
Roughly speaking, the borders of this zoological zone coincide
with the geographical limits of the tropical zone, the tropics of the
Cancer and Capricorn; its characteristic forms, however, extend in
undulating lines several degrees north and southwards.
Commencing from the west coast of Africa the desert of the Sahara
forms a well-marked boundary between the equatorial and northern
zones; as the boundary approaches the Nile it makes a sudden
sweep towards the north as far as Northern Syria (Mastacembelus,
near Aleppo, and in the Tigris; Clarias and Chromides, in the lake of
Galilee); crosses through Persia and Afghanistan (Ophiocephalus),
to the southern ranges of the Himalayas, and follows the course of
the Yang-tse-Kiang, which receives its contingent of equatorial fishes
through its southern tributaries. Its continuation through the North
Pacific may be considered to be indicated by the tropic which strikes
the coast of Mexico at the southern end of the Gulf of California.
Equatorial types of South America are known to extend so far
northwards; and by following the same line the West India Islands
are naturally included in this zone.
Towards the south the equatorial zone embraces the whole of
Africa and Madagascar, and seems to extend still farther south in
Australia, its boundary probably following the southern coast of that
continent; the detailed distribution of the freshwater fishes of South-
Western Australia has been but little studied, but the few facts which
we know show that the tropical fishes of Queensland follow the
principal water-course of that country, the Murray River, far towards
the south and probably to its mouth. The boundary-line then
stretches northwards of Tasmania and New Zealand, coinciding with
the tropic until it strikes the western slope of the Andes, on the South
American Continent, where it again bends southwards to embrace
the system of the Rio de la Plata.
The equatorial zone is divided into four regions:—
A. The Indian region.
B. The African region.
C. The Tropical American region.
D. The Tropical Pacific region.

These four regions diverge into two well-marked divisions, one of

which is characterised by the presence of Cyprinoid fishes,
combined with the development of Labyrinthici; whilst in the other
both these types are absent. The boundary between the Cyprinoid
and Acyprinoid division seems to follow Wallace’s line, a line drawn
from the south of the Philippines between Borneo and Celebes, and
farther south between Bali and Lombock. Borneo abounds in
Cyprinoids; from the Philippine Islands a few only are known at
present, and in Bali two species have been found; but none are
known from Celebes or Lombock, or from islands situated farther
east of them.[20]
Taking into consideration the manner in which Cyprinoids and
Siluroids have been dispersed, we are obliged to place the Indian
region as the first in the order of our treatment; and indeed the
number of its freshwater fishes, which appear to have spread from it
into the neighbouring regions, far exceeds that of the species which
it has received from them.

A. The Indian Region comprises the whole continent of Asia

south of the Himalayas and the Yang-tse-kiang; it includes the
islands to the west of Wallace’s line. Towards the north-east the
island of Formosa, which also by other parts of its fauna leans more
towards the equatorial zone, has received some characteristic
Japanese Freshwater fishes, for instance, the singular Salmonoid
Plecoglossus. Within the geographical boundaries of China the
Freshwater fishes of the tropics pass gradually into those of the
northern zone, both being separated by a broad debateable ground.
The affluents of the great river traversing this district are more
numerous from the south than from the north, and carry the southern
fishes far into the temperate zone. The boundary of this region
towards the north-west is scarcely better defined. Before Persia
passed through the geological changes by which its waters were
converted into brine and finally dried up, it seems to have been
inhabited by many characteristic Indian forms, of which a few still
survive in the tract intervening between Afghanistan and Syria;
Ophiocephalus and Discognathus have each at least one
representative, Macrones has survived in the Tigris, and
Mastacembelus has penetrated as far as Aleppo. Thus, Freshwater
fishes belonging to India, Africa, and Europe, are intermingled in a
district which forms the connecting link between the three continents.
Of the freshwater fishes of Arabia we are perfectly ignorant; so much
only being known that the Indian Discognathus lamta occurs in the
reservoirs of Aden, having, moreover, found its way to the opposite
African coast; and that the ubiquitous Cyprinodonts flourish in
brackish pools of Northern Arabia.
The following is the list of the forms of freshwater fishes
inhabiting this region:[21]—
Percina—
Lates[22] [Africa, Australia] 1 species.
Nandina 7 „
Labyrinthici [Africa] 25 „
Luciocephalidæ 1 „
Ophiocephalidæ [1 species in Africa] 30 „
Mastacembelidæ [3 species in Africa] 10 „
Chromides [Africa, South America]
Etroplus 2 „

Siluridæ—
Clariina [Africa] 12 „
Chacina 3 „
Silurina [Africa, Palæarct.] 72 „
Bagrina [Africa] 50 „
Ariina [Africa, Australia, South America] 40 „
Bagariina 20 „
Rhinoglanina [Africa] 1 „
Hypostomatina [South America] 5 „

Cyprinodontidæ—
Carnivoræ [Palæarct., North America, Africa, South America]
Haplochilus [Africa, South America, North America, Japan] 4 „

Cyprinidæ [Palæarct., N. America, Africa]—

Cyprinina [Palæarct., N. America, Africa] 190 „
Rasborina [Africa, 1 species] 20 „
Semiplotina 4 „
Danionina [Africa] 30 „
Abramidina [Palæarct., N. Amer., Africa] 30 „
Homalopterina 10 „
Cobitidina [Palæarct.] 50 „
Osteoglossidæ [Africa, Australia, S. America] 1 „
Notopteridæ [Africa] 3 „
Symbranchidæ—
Amphipnous 1 „
 Monopterus 1 „
Symbranchus [1 species in S. America] 2 „
625 species.

In analysing this list we find that out of 39 families or groups of

freshwater fishes 12 are represented in this region, and that 625
species are known to occur in it; a number equal to two-sevenths of
the entire number of freshwater fishes known. This large proportion
is principally due to the development of numerous local forms of
Siluroids and Cyprinoids, of which the former show a contingent of
about 200, and the latter of about 330 species. The combined
development of those two families, and their undue preponderance
over the other freshwater types, is therefore the principal
characteristic of the Indian region. The second important character of
its fauna is the apparently total absence of Ganoid and
Cyclostomous fishes. Every other region has representatives of
either Ganoids or Cyclostomes, some of both. However, attention
has been directed to the remarkable coincidence of the geographical
distribution of the Sirenidæ and Osteoglossidæ, and as the latter
family is represented in Sumatra and Borneo, it may be reasonably
expected that a Dipnoous form will be found to accompany it. The
distribution of the Sirenidæ and Osteoglossidæ is as follows:—
Tropical America.
Lepidosiren paradoxa. Osteoglossum bicirrhosum.
Arapaima gigas.

Tropical Australia.
Ceratodus forsteri. Osteoglossum leichardti.
Ceratodus miolepis.

East Indian Archipelago.

? Osteoglossum formosum.

Tropical Africa.
Protopterus annectens. Heterotis niloticus.
Not only are the corresponding species found within the same
region, but also in the same river systems; and although such a
connection may and must be partly due to a similarity of habit, yet
the identity of this singular distribution is so striking that it can only
be accounted for by assuming that the Osteoglossidæ are one of the
earliest Teleosteous types which have been contemporaries of and
have accompanied the present Dipnoi since or even before the
beginning of the tertiary epoch.
Of the autochthont freshwater fishes of the Indian region, some
are still limited to it, viz., the Nandina, the Luciocephalidæ (of which
one species only exists in the Archipelago), of Siluroids the Chacina
and Bagariina, of Cyprinoids the Semiplotina and Homalopterina;
others very nearly so, like the Labyrinthici, Ophiocephalidæ,
Mastacembelidæ, of Siluroids the Silurina, of Cyprinoids the
Rasborina and Danionina, and Symbranchidæ.
The regions with which the Indian has least similarity are the
North American and Antarctic, as they are the most distant. Its
affinity to the other regions is of a very different degree:—
1. Its affinity to the Europo-Asiatic region is indicated almost
solely by three groups of Cyprinoids, viz., the Cyprinina, Abramidina,
and Cobitidina. The development of these groups north and south of
the Himalayas is due to their common origin in the highlands of Asia;
but the forms which descended into the tropical climate of the south
are now so distinct from their northern brethren that most of them are
referred to distinct genera. The genera which are still common to
both regions are only the true Barbels (Barbus), a genus which, of all
Cyprinoids, has the largest range over the old world, and of which
some 160 species have been described; and, secondly, the
Mountain Barbels (Schizothorax, etc.), which, peculiar to the Alpine
waters of Central Asia, descend a short distance only towards the
tropical plains, but extend farther into rivers within the northern
temperate districts. The origin and the laws of the distribution of the
Cobitidina appear to have been identical with those of Barbus, but
they have not spread into Africa.
If, in determining the degree of affinity between two regions, we
take into consideration the extent in which an exchange has taken
place of the faunæ originally peculiar to each, we must estimate that
obtaining between the freshwater fishes of the Europo-Asiatic and
Indian regions as very slight indeed.
 2. There exists a great affinity between the Indian and African
regions; seventeen out of the twenty-six families or groups found in
the former are represented by one or more species in Africa, and
many of the African species are not even generically different from
the Indian. As the majority of these groups have many more
representatives in India than in Africa, we may reasonably assume
that the African species have been derived from the Indian stock; but
this is probably not the case with the Siluroid group of Clariina, which
with regard to species is nearly equally distributed between the two
regions, the African species being referable to three genera (Clarias,
Heterobranchus, Gymnallabes, with the subgenus Channallabes),
whilst the Indian species belong to two genera only, viz. Clarias and
Heterobranchus. On the other hand, the Indian region has derived
from Africa one freshwater form only, viz. Etroplus, a member of the
family of Chromides, so well represented in tropical Africa and South
America. Etroplus inhabits Southern and Western India and Ceylon,
and has its nearest ally in a Madegasse Freshwater fish,
Paretroplus. Considering that other African Chromides have
acclimatised themselves at the present day in saline water, we think
it more probable that Etroplus should have found its way to India
through the ocean than over the connecting land area; where,
besides, it does not occur.
3. A closer affinity between the Indian and Tropical American
regions than is indicated by the character of the equatorial zone
generally, does not exist. No genus of Freshwater fishes occurs in
India and South America without being found in the intermediate
African region, with two exceptions. Four small Indian Siluroids
(Sisor, Erethistes, Pseudecheneis, and Exostoma) have been
referred to the South American Hypostomatina; but it remains to be
seen whether this combination is based upon a sufficient agreement
of their internal structure, or whether it is not rather artificial. On the
other hand, the occurrence and wide distribution in tropical America
of a fish of the Indian family Symbranchidæ (Symbranchus
marmoratus), which is not only congeneric with, but also most
closely allied to, the Indian Symbranchus bengalensis, offers one of
those extraordinary anomalies in the distribution of animals of which
no satisfactory explanation can be given at present.
 4. The relation of the Indian region to the Tropical Pacific region
consists only in its having contributed a few species to the poor
fauna of the latter. This immigration must have taken place within a
recent period, because some species now inhabit fresh waters of
tropical Australia and the South Sea Islands without having in any
way changed their specific characters, as Lates calcarifer, species of
Dules, Plotosus anguillaris; others (species of Arius) are but little
different from Indian congeners. All these fishes must have migrated
by the sea; a supposition which is supported by what we know of
their habits. We need not add that India has not received a single
addition to its freshwater fish-fauna from the Pacific region.
Before concluding these remarks on the Indian region, we must
mention that peculiar genera of Cyprinoids and Siluroids inhabit the
streams and lakes of its alpine ranges in the north. Some of them,
like the Siluroid genera Glyptosternum, Euglyptosternum,
Pseudecheneis, have a folded disk on the thorax between their
horizontally spread pectoral fins; by means of this they adhere to
stones at the bottom of the mountain torrents, and without it they
would be swept away into the lower courses of the rivers. The
Cyprinoid genera inhabiting similar localities, and the lakes into
which the alpine rivers pass, such as Oreinus, Schizothorax,
Ptychobarbus, Schizopygopsis, Diptychus, Gymnocypris, are
distinguished by peculiarly enlarged scales near the vent, the
physiological use of which has not yet been ascertained. These
alpine genera extend far into the Europo-Asiatic region, where the
climate is similar to that of their southern home. No observations
have been made by which the altitudinal limits of fish life in the
Himalayas can be fixed, but it is probable that it reaches the line of
perpetual snow, as in the European Alps which are inhabited by
Salmonoids. Griffith found an Oreinus and a Loach, the former in
abundance, in the Helmund at Gridun Dewar, altitude 10,500 feet;
and another Loach at Kaloo at 11,000 feet.
 B. The African Region comprises the whole of the African
continent south of the Atlas and the Sahara. It might have been
conjectured that the more temperate climate of its southern extremity
would have been accompanied by a conspicuous difference of the
fish-fauna. But this is not the case; the difference between the
tropical and southern parts of Africa consists simply in the gradual
disappearance of specifically tropical forms, whilst Siluroids,
Cyprinoids, and even Labyrinthici penetrate to its southern coast; no
new form has entered to impart to South Africa a character distinct
from the central portion of the continent. In the north-east the African
fauna passes the Isthmus of Suez and penetrates into Syria; the
system of the Jordan presenting so many African types that it has to
be included in a description of the African region as well as of the
Europo-Asiatic. This river is inhabited by three species of Chromis,
one of Hemichromis, and Clarias macracanthus, a common fish of
the Upper Nile. This infusion of African forms cannot be accounted
for by any one of those accidental means of dispersal, as
Hemichromis is not represented in the north-eastern parts of Africa
proper, but chiefly on the west coast and in the Central African lakes.
Madagascar clearly belongs to this region. Besides some Gobies
and Dules, which are not true freshwater fishes, four Chromides are
known. To judge from general accounts, its Freshwater fauna is
poorer than might be expected; but, singular as it may appear,
collectors have hitherto paid but little attention to the Freshwater
fishes of this island. The fishes found in the freshwaters of the
Seychelles and Mascarenes are brackish-water fishes, such as
Fundulus, Haplochilus, Elops, Mugil, etc.
The following is the list of the forms of Freshwater fishes
inhabiting this region:—
Dipnoi [Australia, Neotrop.]—
Lepidosiren annectens 1 species.
Polypteridæ 2 „

Percina (Cosmopol.)—
Lates [India, Australia] 1 „
Labyrinthici [India] 5 „
Ophiocephalidæ [India] 1 „
Mastacembelidæ [India] 3 „

Chromides [South America]—

Chromis 23 „
Hemichromis 5 „
Paretroplus 1 „

Siluridæ—
Clariina [India] 14 „
Silurina [India, Palæarct.] 11 „
Bagrina [India] 10 „
Pimelodina [South America] 2 „
Ariina[23] [India, Australia, S. Amer., Patagonia] 4 „
Doradina [South America]—
Synodontis 15 „
Rhinoglanina [India] 2 „
Malapterurina 3 „

Characinidæ [South America]—

Citharinina 2 „
Nannocharacina 2 „
Tetragonopterina—
Alestes 14 „
Crenuchina—
Xenocharax 1 „
Hydrocyonina—
Hydrocyon 4 „
Distichodontina 10 „
Ichthyborina 2 „
Mormyridæ (Gymnarchidæ) 51 „

Cyprinodontidæ—
Carnivoræ [Palæarct., India, S. America—
Haplochilus [India, South America] 7 „
Fundulus [Palæarct., Nearct.] 1 „

Cyprinidæ [Palæarct., India, North America]—

Cyprinina [Palæarct., India, N. America—
Labeo [India] 6 „
Barynotus [India] 2 „
Abrostomus 2 „
Discognathus lamta[24] [India] 1 „
Barbus [Palæarct., India] 35 „