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Generalship in Ancient Greece Rome and Byzantium 1st
Edition Shaun Tougher Digital Instant Download
Author(s): Shaun Tougher, Richard Evans
ISBN(s): 9781474459969, 147445996X
Edition: 1
File Details: PDF, 19.21 MB
Year: 2022
Language: english
Generalship in Ancient Greece,
Rome and Byzantium
For NCT
Generalship in Ancient Greece,
Rome and Byzantium
Edited by Richard Evans and Shaun Tougher
 Edinburgh University Press is one of the leading university presses in the UK. We publish
academic books and journals in our selected subject areas across the humanities and social
sciences, combining cutting-edge scholarship with high editorial and production values
to produce academic works of lasting importance. For more information visit our website:
edinburghuniversitypress.com

© editorial matter and organisation Richard Evans and Shaun Tougher, 2022
© the chapters their several authors, 2022

Cover image: Tapestry showing Constantine’s Triumphal Entry into Rome 1623–1625.
Figural composition designed in 1622 by Peter Paul Rubens. Philadelphia Museum
of Art: Gift of the Samuel H. Kress Foundation, 1959, 1959-78-1.
Cover design: Workhaus.

Edinburgh University Press Ltd

The Tun­– H
­ olyrood Road, 12(2f) Jackson’s Entry, Edinburgh EH8 8PJ

Typeset in 10/14 Ehrhardt

by Cheshire Typesetting Ltd, Cuddington, Cheshire, and
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A CIP record for this book is available from the British Library

ISBN 978 1 4744 5994 5 (hardback)

ISBN 978 1 4744 5996 9 (webready PDF)
ISBN 978 1 4744 5997 6 (epub)

The right of Richard Evans and Shaun Tougher to be identified as the editors of this work has
been asserted in accordance with the Copyright, Designs and Patents Act 1988, and the Copyright
and Related Rights Regulations 2003 (SI No. 2498).
Contents

Acknowledgements vii
List of Contributors ix
Abbreviations xii

Introduction1
Richard Evans and Shaun Tougher

1. Kings, Tyrants and Bandy-Legged Men: Generalship in Archaic

Greece6
Cezary Kucewicz

2. Commemorating Thermopylae: The andreia of Glorious Defeat

as a Literary Construct 36
Richard Evans

3. Plato on Military and Political Leadership 52

Nicholas Rockwell

4. Reconstructing Early Seleucid Generalship, 301–222 bc67

Alex McAuley

5. Generalship and Knowledge in the Middle Roman Republic 86

Michael Taylor

6. Command Assessment in the Bellum Gallicum: Caesar and Fortuna98

David Nolan
vi | CONTENTS

7. Remembering P. Quinctilius Varus: Opposing Perspectives on the

Memory and Memorialisation of the Failed General in the Annales
of Tacitus116
Daniel Crosby

8. Decius and the Battle near Abritus 139

David Potter

9. Ammianus and the Heroic Mode of Generalship in the Fourth

Century ad151
Conor Whately

10. The Fine Line between Courage and Fear in the Vandal War164
Michael Stewart

11. The Generalship of John Troglita: Art in Artifice 187

Martine de Marre

12. The Best of Men: Cross-Cultural Command in the 630s ad206

Eve MacDonald

13. Tian Yue Marshals His Tropes: Public Persuasion and the Character of
Military Leadership in Late Tang China 225
David A. Graff

14. The Ideal of the Roman General in Byzantium: The Reception of

Onasander’s Strategikos in Byzantine Military Literature 242
Philip Rance

15. Generalship and Gender in Byzantium: Non-Campaigning Emperors

and Eunuch Generals in the Age of the Macedonian Dynasty 264
Shaun Tougher

16. The Politics of War: Virtue, Tyche, Persuasion and the Byzantine
General284
Dimitris Krallis

Epilogue306
Richard Evans and Shaun Tougher

Bibliography 307
Index 351
Acknowledgements

This volume has its origins in a panel on ‘The Art of Generalship: Late Antique,
Byzantine, and Chinese Ideals’ held at the International Medieval Congress in Leeds
in 2014. For this publication the scope of the original panel was extended to encom-
pass the ancient Graeco-Roman world. In the course of its production several debts
of gratitude have been incurred. We would like to thank all the contributors to the
original panel, including those who were not able to contribute to the final volume:
Doug Lee, Peter Lorge, Ken Swope and Jamie Wood. We are also indebted to the
many colleagues around the world who acted as readers for the individual chap-
ters and supplied helpful and constructive feedback. In addition, we cannot thank
enough Edinburgh University Press for their support and extreme patience, espe-
cially Carol Macdonald. The schedule faced of course not just the usual obstructions
to finalisation experienced by academics but also the ongoing Covid pandemic.
Finally, Shaun would like to thank Richard for his immense contribution to the
completion of the volume; he proved as excellent and positive a co-editor as he was
a colleague at Cardiff.

Richard Evans and Shaun Tougher

Pretoria and Cardiff
List of Contributors

Daniel Crosby recently completed his PhD studies at Bryn Mawr College,
Philadelphia, USA. While the main focus of much of his forthcoming and ongoing
research is on divination in the ancient world, he has also published on Roman
prosopography (2016), on Patristics (2017) and in this volume on Tacitus and the
dynamics of cultural memory.

Martine de Marre is currently Associate Professor of Ancient History in the

Department of Biblical and Ancient Studies at the University of South Africa,
Pretoria. Her research focuses on the socio-cultural history of Roman North Africa,
especially the role of women, and the works of Augustine, Fulgentius and notably
Corippus (2020).

Richard Evans taught at the University of South Africa, Pretoria, and Cardiff
University, Wales. He is currently an Academic Associate in the Department of
Biblical and Ancient Studies, University of South Africa. He has authored books
on Gaius Marius (1994), the Roman Republic (2003), ancient Syracuse (2009 and
2016), Pergamum (2012) and ancient warfare (2010, 2013 and 2015). He has also
edited or co-edited volumes on mass and elite interaction in antiquity (2017), ancient
divination (2018) and piracy in ancient Greece and Rome (2020).

David A. Graff is Professor in the Department of History at Kansas State University,

Manhattan, USA. Since 2017 he has held the Richard A. and Greta Bauer Pickett
Chair for Exceptional Faculty. He has published widely on Chinese military history,
with a particular focus on the Tang Dynasty (2002) and the comparative military
practice of China and Byzantium (2016).
x | LIST OF CONTRIBUTORS

Dimitris Krallis is Professor at the Department of Humanities and Director of

the Stavros Niarchos Foundation Centre for Hellenic Studies at Simon Fraser
University, Canada. He has published on the social, political and intellectual his-
tory of the Byzantine Empire, with books on the life, work and career of Michael
Attaleiates (2012 and 2019).

Cezary Kucewicz completed an MA in Ancient History at Cardiff University,

Wales, and more recently his PhD at University College London, England, on
Homeric warfare. He is currently a postdoctoral fellow at the University of Gdańsk,
Poland, and Wolfson College, Cambridge, England. He is the author of The
Treatment of the War Dead in Archaic Athens: An Ancestral Custom (2021).

Alex McAuley is Senior Lecturer in Hellenistic History at Cardiff University,

Wales. He has published extensively on the Hellenistic period, notably on the accul-
turation process between Macedonians and Mesopotamian groups at the start of
Seleucid rule. Royal women in the dynastic structures of the Diadochi have also been
a focus of recent research.

Eve MacDonald is a Lecturer in Ancient History at Cardiff University, Wales.

She is interested in the social and military history of the Sasanian period and its
connections to the Byzantine and early Islamic worlds. She has published on the
archaeology of the Sasanian frontiers, and on Carthage and North Africa.

David Nolan completed his PhD at the University of Tasmania, Australia. His
research interests focus on Caesar’s leadership in his conquest of Gaul, on the battle
narrative in the Bellum Gallicum and on the role of the centurions (2016).

David Potter is Professor of Ancient History at the University of Michigan, Ann

Arbor, USA. His research interests encompass Greek and Latin historiography,
public entertainments in the Graeco-Roman world, ancient warfare, and Greek and
Roman Asia Minor. His publications are many, including Emperors of Rome (2008),
The Roman Empire at Bay (2014) and The Origin of Empire (2018).

Philip Rance obtained his PhD at St Andrews, Scotland. He has taught History
and Greek Language at universities in the UK and Germany, and held numerous
research fellowships in Europe. He is a Visiting Scholar at the Free University,
Berlin, Germany, and a Research Fellow at the Centre for Advanced Study, Sofia,
Bulgaria. He has published extensively on late antique and Byzantine history and
literature.
 LIST OF CONTRIBUTORS | xi

Nicholas Rockwell is a Lecturer in Political Science and History at the University

of Colorado, Denver, USA. He obtained his PhD from the University of California,
Los Angeles, USA, and his MA and BA from California State University, Fresno,
USA. He is the author of a monograph entitled Thebes: A History (2017), and is cur-
rently engaged in research towards a study of the citizen-soldier in the ancient world.

Michael Stewart is an Honorary Research Fellow in the School of History and

Philosophical Inquiry at the University of Queensland, Australia. Having completed
his PhD thesis in 2013 he has since published extensively, including The Soldier’s
Life: Martial Virtues and Manly Romanitas in the Early Byzantine Empire (2016).

Michael Taylor is an Assistant Professor in the History Department, State

University of New York at Albany, USA. He obtained his PhD from the University
of California, Berkeley, USA. He has published widely on Roman Republican war-
fare and political life, and is the author of Soldiers and Silver: Mobilizing Resources in
the Age of Roman Conquest (2020).

Shaun Tougher is Professor of Late Roman and Byzantine History at Cardiff

University, Wales. His numerous publications include The Reign of Leo VI (1997),
Julian the Apostate (2007), The Eunuch in Byzantine History and Society (2008) and,
most recently, The Roman Castrati: Eunuchs in the Roman Empire (2021).

Conor Whately is Associate Professor in Classics at the University of Winnipeg,

Canada. He has written or co-edited six books. One was a co-edited volume with
James Chlup on Greek and Roman military manuals (2020), one was on the Roman
military in the imperial-era Balkans (2016), and two were on assorted aspects of
Procopius’ writing as they pertain to war and war-making (2016 and 2021). Two
books are for a more general audience, one an introduction to the Roman military
(2020), the other an introduction to sensory aspects of ancient warfare (2021).
Abbreviations

The following is a list of abbreviations of key journals and collections listed in the
bibliography. Ancient writers and their works are generally abbreviated according to
current conventions, although some contributors adopt variations. Likewise, indi-
vidual contributors follow personal preference in the spelling of names and trans-
literation of ancient Greek. Regarding dating abbreviations, bc and ad are adopted
throughout the volume.

AClass Acta Classica

AFLS Annali della Facoltà di Lettere e Filosofia dell’Università di Siena
AHB The Ancient History Bulletin
AJP American Journal of Philology
Anc. Soc. Ancient Society
AnnBari Annali della Facoltà di Lettere e Filosofia, Università di Bari
ANRW Aufstieg und Niedergang der römischen Welt
Ant. Class. L’Antiquité Classique
BCH Bulletin de Correspondence Hellénique
BF Byzantinische Forschungen
BMGS Byzantine and Modern Greek Studies
Byz Byzantion
BZ Byzantinische Zeitschrift
C&M Classica et Mediaevalia
CJ Classical Journal
Cl. Ant. Classical Antiquity
CP Classical Philology
CQ Classical Quarterly
 ABBREVIATIONS | xiii

CR Classical Review
CW Classical World
DHA Dialogues d’Histoire Ancienne
DOP Dumbarton Oaks Papers
EA Epigraphica Anatolica
G&R Greece and Rome
GRBS Greek, Roman and Byzantine Studies
Hist. Historia
JAH Journal of Ancient History
JHS Journal of Hellenic Studies
JJS Journal of Jewish Studies
JLA Journal of Late Antiquity
JÖB Jahrbuch der Österreichischen Byzantinistik
JRA Journal of Roman Archaeology
JRS Journal of Roman Studies
MAAR Memoirs of the American Academy in Rome
MD Materiali e Discussioni
MH Museum Helveticum
NC The Numismatic Chronicle
P&P Past and Present
PBSR Papers of the British School at Rome
PG Patrologia Graeca
PhW Philologische Wochenschrift
PLLS Papers of the Leeds Latin Seminar
RE Paulys Real-Encyclopädie der classischen Altertumswissenschaft
REA Revue des Études Anciennes
Rev. Ét. Grec. Revue des Études Grecques
RhM Rheinisches Museum
TAPA Transactions of the American Philological Association
TM Travaux et Mémoires
WJA Würzburger Jahrbücher für die Altertumswissenschaft
ZPE Zeitschrift für Papyrologie und Epigraphik
 Introduction
Richard Evans and Shaun Tougher

’Tis not in mortals to command success,

But we’ll do more, Sempronius; we’ll deserve it.
Joseph Addison, Cato: A Tragedy, 1.2.43

The art of generalship or the ability of successful leadership in a military context, as

Addison perceptively observed, is not at all dependent on fortune or tyche (meaning
in reality good luck) to ensure victory. Victory might be obtained through good luck
just happening,1 but the soundest method of ensuring success relies on fastidious
management in the camp and while on the march in conjunction with inspiring
leadership on the field of battle. How many or how few generals in antiquity measure
up to such a stringent assessment of their capabilities, their practice of the art of
being a general, and so deserved the successes they obtained and their subsequent
immortal fame? In the following chapters the discussion aligns to this focus. Julius
Caesar, for example, makes frequent reference to ‘good fortune’ in his Gallic cam-
paigns (Nolan), but he was an able if not the consummate manager of his military
adventures, in Gaul, Italy, Spain and Greece in the first century bc.2 In political life
he perhaps lacked that lightness of touch which his successor Augustus, only a mod-
erately capable military leader, possessed in abundance. Leonidas, the Spartan king

1
Such as a timely thunderstorm, a dust storm or a snowstorm, or a fortuitous eclipse of the
sun or moon. All these natural phenomena could affect the moods of the opposing sides in
military campaigns, but could also be explained as divine intervention.
2
His expeditions to Britain in 55 and 54 bc, however, showed the limitations of ancient
access to information and the logistical difficulties relating to organising long-range sea-
borne expeditions.
2 | RICHARD EVANS AND SHAUN TOUGHER

(Evans), and the Roman emperor Decius (Potter), on the other hand, no doubt both
capable soldiers, found immortality not in success, but in two of the most famous
defeats in the history of the Greeks and Romans: Thermopylae in 480 bc and Abritus
in ad 251 respectively.
The importance to cultures in antiquity of success in military affairs can easily
be measured by noting that Homer’s Iliad, among the earliest surviving literary
evidence for the history and civilisation of the ancient Greeks, is essentially a tale
in verse about the flawed leadership of the protagonists: Agamemnon, Achilles and
Hector. All are active participants in the military campaigning around Troy and the
battles that took place there (Kucewicz). None is the perfect general, but Alexander
the Great (336–323 bc), probably the greatest warrior general of history, and the
most sublime practitioner of generalship, constantly sought to emulate these myth-
ical figures; and he was not an isolated example. On the other hand, the Roman
emperor Trajan (ad 98–117), certainly regarded by the Romans as completely equal
in ability to Alexander, was, very much like Caesar, a manager of his armies and not
a front-line fighter. In late antiquity, the ambitious eastern campaign of emperor
Julian (361–363) can be presented or even understood as a desire to follow in the
footsteps of Alexander the Great and the Homeric heroes. Julian’s death on the field
of battle on 26 June 363 was in keeping with the by then thousand-year-old tradition
of the Graeco-Roman warrior.3 Further, he was certainly not the last, as the follow-
ing discussions will illustrate. Indeed, Julian’s words are as revealing as his actions.
Well known for his writings as much as his political and military career, in his cele-
brated satire The Caesars – in which the gods undertake to decide who has been the
best Roman emperor of all – three of the six finalists were chosen as being the most
warlike: Julius Caesar, Augustus and Trajan.4 Further, although Marcus Aurelius
was selected as a philosopher,5 and Constantine was thrown into the mix as a lover of
pleasure,6 both emperors also played a military role, and as much as Julian sought to
downplay it, Constantine was a highly successful general. The one Greek figure who
was allowed to be included in the competition and as one of the finalists reflects the
prevalence of the military role of leaders too, for it was Alexander the Great.7
Homeric generalship involved conspicuous participation on the battlefield
and could and did result in some of the most memorable leadership qualities dis-
played throughout antiquity by, for example, not only Alexander the Great, the

3
Amm. Marc. 25.3.
4
Caesars 317B.
5
Caesars 317C.
6
Caesars 317D–318A.
7
Caesars 316B–D.
 INTRODUCTION | 3

early Seleucids (McAuley) or Julius Caesar, but also Epaminondas of Thebes (died
362 bc), Agesilaus of Sparta (c. 400–360 bc) and Dionysius I of Syracuse (405–
367 bc). It could also result in disaster, not only for Julian but also, for example,
for Antigonus Monophthalmus or Pyrrhus of Epirus, both killed fighting at vital
times for the fortunes of their states, in 301 bc and 272 bc respectively. Homeric
generalship also belonged to an epoch in which the leader of a state was first and
perhaps only a warrior, or portrayed as such: Odysseus, Ajax, even Nestor (when he
was a young man). But the increasing sophistication of states and how they operated
in the ancient world brought with it a change in the way military governance was
conceived. Civic leadership, which required competencies across a number of vital
fields from the juridical to oratorical ability for meetings of citizen assemblies, and
civil administration, brought with it a diminution of the purely military career. By
the fifth century bc in Athens leaders such as Miltiades, Aristides, Themistocles,
Pericles and Nicias possessed as many strengths as administrators as they did as
generals. Their successes in peace as in war indicate not only a wealth of talent
in Athenian society, but also that those who reached the pinnacle of success were
extraordinary all-rounders. From the fifth century on, there were as many gener-
als who directed their armies from behind the front line of fighting as there were
Homeric-type warrior generals on the battlefield. While in the first half of the fourth
century bc the Theban leaders Pelopidas and Epaminondas were likely to face oppo-
nents such as Agesilaus of Sparta, Alexander of Pherae or Iphicrates of Athens on the
front line, there were equally successful generals who saw little or no actual combat.8
Philip II (359–336 bc), the Macedonian monarch who was father of a greater son,
was highly successful in pursuit of territorial acquisitions, but he is not remembered
in the same way as his warrior-king son Alexander the Great.9
The expansion into different areas of concern and the growing complexities of
management in military affairs must have been the primary cause for writers such
as Aeneas Tacticus and Xenophon in the fourth century bc to circulate reflections
on how campaigns might be organised by a general, the sort of training needed to
become a good commander, and the various elements of campaigns: from battlefield
encounters to armies on the march, and even the preparations and techniques need
to besiege or defend a city. It is evident that both the specific problems confronting

8
Pelopidas was killed on the battlefield. Epaminondas and Agesilaus were active in combat
well into old age. Epaminondas died at the Battle of Mantinea in 372 bc, Agesilaus while
returning from serving as a mercenary in Egypt in 350 bc.
9
Philip lost the sight of an eye at the siege of Methone, but he appears to have been at some
distance from the front line when he suffered this injury. At his greatest victory, Chaeronea
in 338 bc, Philip commanded the right wing but it was his son on the left wing who is
recorded as being in the thick of the fighting.
4 | RICHARD EVANS AND SHAUN TOUGHER

generals and the wider issues of what was implicit in generalship or leadership began
to be a topic of debate among intellectual circles, as can easily be gauged from the
extant works of Plato and Aristotle (Rockwell), and remained so even a full millen-
nium later in the Byzantine Empire, as reflected by such texts at the Strategikon
ascribed to the emperor Maurice (582–602) and the Taktika of the emperor Leo VI
(886–912) (Rance, Tougher, Krallis).
As yet there was no specialised military elite as is found today in the gradu-
ates of numerous military academies scattered around the world. The general was
very much an amateur, as exemplified by the commanders of armies in the Roman
Republic (Taylor), and indeed the phenomenon persisted until the First World War
(1914–1918). As such these amateurs could, and often did, scale dizzy heights in
their successes: Scipio Africanus at Zama in 202 bc, the younger Aemilius Paullus at
Pydna in 168 bc, Gaius Marius at Vercellae in 101 bc; but they might also plumb the
depths of catastrophic disasters: Gaius Flaminius at Lake Trasimene in 217 bc, the
elder Aemilius Paullus at Cannae in 216 bc, or Augustus’ general Quinctilius Varus
at the Teutoburg Wald in ad 9 (Crosby).
By the time of the early Roman Empire, the rulers, although commanders-in-chief
of their armies stationed on the far-flung frontiers, rarely ventured from the metrop-
olis. Active generalship roles were therefore the occupation of subordinates, but
victories, and by implication defeats,10 belonged to the ruler and reflected either his
glory or his lack of talent. By the mid-second century ad Roman emperors had taken
to leading campaigns personally if not actually fighting in battles. Marcus Aurelius
(161–180) spent much of his rule directing efforts to prevent Germanic tribes from
penetrating the Roman Empire’s northern borders. From the rule of Septimius
Severus (193–211) onwards it again became almost a rule of thumb that the emperor
was active in campaigns. Decius (249–251) was the first Roman emperor to die in
battle against invaders. Further, the third century was the age of the soldier emper-
ors, the phenomenon that many of those who became emperor had risen through the
ranks of the army. This began with Maximinus Thrax (235–238) but is exemplified
by the famous figure of Diocletian (284–305). This age of emperor generals included
Constantine the Great (306–337), Constantius II (337–361), Julian (355–363),
Valentinian I (364–375) and his brother Valens (364–378) (Whately), but the death
of Theodosius I (379–395) marked a return to the figure of the non-campaigning
emperor. This shift began with Theodosius’ young sons and successors Arcadius
(395–408) and Honorius (395–423), but is famously exemplified by Justinian I
(527–565), who nevertheless presided over the reconquest of North Africa and Italy,

10
Victories were often celebrated lavishly, while defeats could be blamed on the commander
in the field rather than on the emperor.
Other documents randomly have
different content
 We need not follow Darwin through his discussion of those cases
in which the adults have a winter and a summer dress and the
young resemble the one or the other in plumage, or are different
from either. The discussion of these cases, confessedly very
complex, adds nothing to our understanding of the theory, and little
but conjecture is offered to account for the facts.
The extreme to which even conjecture can be carried may be
gathered from the following quotation, taken from the section
dealing with cases in which the young in their first plumage differ
from each other according to sex, the young males resembling more
or less closely the adult males, and the young females more or less
closely the adult females:
“Two humming-birds belonging to the genus Eustephanus, both
beautifully colored, inhabit the small island of Juan Fernandez, and
have always been ranked as specifically distinct. But it has lately
been ascertained that the one which is of a rich chestnut-brown
color with a golden-red head, is the male, whilst the other, which is
elegantly variegated with green and white with a metallic-green
head, is the female. Now the young from the first somewhat
resemble the adults of the corresponding sex, the resemblance
gradually becoming more and more complete.
“In considering this last case, if as before we take the plumage of
the young as our guide, it would appear that both sexes have been
rendered beautiful independently; and not that one sex has partially
transferred its beauty to the other. The male apparently has acquired
his bright colors through sexual selection in the same manner as, for
instance, the peacock or pheasant in our first class of cases; and the
female in the same manner as the female Rhynchæa or Turnix in our
second class of cases. But there is much difficulty in understanding
how this could have been effected at the same time with the two
sexes of the same species. Mr. Salvin states, as we have seen in the
eighth chapter, that with certain humming-birds the males greatly
exceed the females in number, whilst with other species inhabiting
the same country the females greatly exceed the males. If, then, we
might assume that during some former lengthened period the males
of the Juan Fernandez species had greatly exceeded the females in
number, but that during another lengthened period the females had
far exceeded the males, we could understand how the males at one
time, and the females at another, might have been rendered
beautiful by the selection of the brighter-colored individuals of either
sex; both sexes transmitting their characters to their young at a
rather earlier age than usual. Whether this is the true explanation I
will not pretend to say; but the case is too remarkable to be passed
over without notice.”
The third group of cases include those in which the females are
more brightly colored, or more ornamented, than the males. These
cases are rare, and the differences between the sexes are never so
great as when the male is the more highly colored. Wallace thinks
that since in these cases the male incubates the eggs his less
conspicuous colors have been acquired through natural selection. In
the genus Turnix the female is larger than the male, and lacks the
black on the throat and neck, and the plumage as a whole is lighter
than that of the male. The natives assert that the females after
laying their eggs associate in flocks, and leave the males to do the
incubating; and from other evidence Darwin thinks that this is true.
In three species of painted snipe the females “are not only larger
but much more richly colored than the males,” and the trachea is
more convoluted in some species. “There is also reason to believe
that the male undertakes the duty of incubation.” In the dotterel
plover the female is larger and somewhat more strongly colored. The
males take at least a share in the incubation. In the common
cassowary the female is larger and the skin of the head more
brightly colored than in the male. The female is pugnacious during
the breeding season and the male sits on the eggs. The female emu
is large and has a crest. She is more courageous and pugilistic and
makes a deep, hollow, guttural boom. The male is more docile and
can only hiss or croak. He not only incubates the eggs, but defends
the young against their own mother. “So that with this emu we have
a complete reversal not only of the parental and incubating instincts,
but of the usual moral qualities of the two sexes; the females being
savage, quarrelsome, and noisy, the males gentle and good. The
case is very different with the African ostrich, for the male is
somewhat larger than the female and has finer plumes with more
strongly contrasted colors; nevertheless he undertakes the whole
duty of incubation.”
Darwin attempts to explain these reversals of instincts on the
assumption that the males have turned the tables on the females,
and have themselves done the selecting; and incidentally, it may be
pointed out in passing, they have had to pay the penalty by
incubating the eggs.
In the group of mammals, Darwin thinks that the male wins the
female by conquering other males rather than by charming her
through his display. The males, even when unarmed, engage in
desperate conflicts with each other, and sometimes kill, but more
often only wound, their fellows. The secondary sexual characters of
the males have been acquired, therefore, by natural selection
applied to one sex, and less frequently through the choice of the
female. Since we are here more especially concerned with the latter
class of phenomena, we may examine only a few cases under the
first head.
The horns of stags are used by them in their conflicts with each
other; the tusks of the elephant make this animal the most
dangerous in the world, when in must. The horns of bulls, the
canine teeth of many mammals, the tusks of the walrus, are further
examples of organs which have been, according to Darwin, acquired
through the competitions of the males with each other.
The voices of mammals are used for various purposes, “as a signal
of danger, as a call from one member of the troup to another, and
from the mother to her lost offspring, or from the latter for
protection.”
“Almost all male animals use their voices much more during the
rutting season than at any other time; and some, as the giraffe and
porcupine, are said to be completely mute excepting at this season.
As the throats (i.e. the larynx and thyroid bodies) of stags
periodically become enlarged at the beginning of the breeding
season, it might be thought that their powerful voices must be
somehow of high importance to them; but this is very doubtful.
From information given to me by two experienced observers, Mr.
McNeill and Sir P. Egerton, it seems that young stags under three
years old do not roar or bellow; and that the old ones begin
bellowing at the commencement of the breeding season, at first only
occasionally and moderately, whilst they restlessly wander about in
search of the females. Their battles are prefaced by loud and
prolonged bellowing, but during the actual conflict they are silent.
Animals of all kinds which habitually use their voices utter various
noises under any strong emotion, as when enraged and preparing to
fight; but this may merely be the result of nervous excitement,
which leads to the spasmodic contraction of almost all the muscles
of the body, as when a man grinds his teeth and clenches his fists in
rage or agony. No doubt stags challenge each other to mortal
combat by bellowing; but those with the more powerful voices,
unless at the same time the stronger, better-armed, and more
courageous, would not gain any advantage over their rivals.”
“Some writers suggest that the bellowing serves as a call to the
female; but the experienced observers above quoted inform me that
female deer do not search for the male, though the males search
eagerly for the females, as indeed might be expected from what we
know of the habits of other male quadrupeds. The voice of the
female, on the other hand, quickly brings to her one or more stags,
as is well known to the hunters who in wild countries imitate her cry.
“As the case stands, the loud voice of the stag during the breeding
season does not seem to be of any special service to him, either
during his courtship or battles, or in any other way. But may we not
believe that the frequent use of the voice, under the strong
excitement of love, jealousy, and rage, continued during many
generations, may at last have produced an inherited effect on the
vocal organs of the stag, as well as of other male animals? This
appears to me, in our present state of knowledge, the most probable
view.”
Here once more we find that Darwin makes use, as a sort of last
resort, of the principle of the inheritance of acquired characters. As
long as the theory of selection, in any of its forms, appears to offer a
satisfactory solution, we find the facts used in support of this theory,
but as soon as a difficulty arises the Lamarckian theory is brought to
the front. It is this shifting, as we have already more than once
pointed out, that shows how little real basis there is for the theory of
sexual selection.
The male gorilla has a tremendous voice, and he has, as has also
the orang, a laryngeal sac. One species of gibbon has the power of
producing a correct octave of musical notes.
“The vocal organs of the American Mycetes caraya are one-third
larger in the male than in the female, and are wonderfully powerful.
These monkeys in warm weather make the forests resound at
morning and evening with their overwhelming voices. The males
begin the dreadful concert, and often continue it during many hours,
the females sometimes joining in with their less powerful voices. An
excellent observer, Rengger, could not perceive that they were
excited to begin by any special cause; he thinks that, like many
birds, they delight in their own music, and try to excel each other.
Whether most of the foregoing monkeys have acquired their
powerful voices in order to beat their rivals and charm the females—
or whether the vocal organs have been strengthened and enlarged
through the inherited effects of long-continued use without any
particular good being thus gained—I will not pretend to say; but the
former view, at least in the case of the Hylobates agilis, seems the
most probable.”
The odor of some mammals is confined to, or more developed, in
the males; but in some forms, as in the skunk, it is present in both
sexes. In the shrew mice, abdominal scent glands are present, but
since these mice are rejected by birds of prey, their glands probably
serve to protect them; “nevertheless the glands become enlarged in
the males during the breeding season.” In many other quadrupeds
the scent glands are of the same size in both sexes, and their
function is unknown.
“In other species the glands are confined to the males, or are
more developed than in the females; and they almost always
become more active during the rutting season. At this period the
glands on the sides of the face of the male elephant enlarge, and
emit a secretion having a strong musky odor. The males, and rarely
the females, of many kinds of bats have glands and protrudable sacs
situated in various parts; and it is believed that these are
odoriferous.
“The rank effluvium of the male goat is well known, and that of
certain male deer is wonderfully strong and persistent. Besides the
general odor, permeating the whole body of certain ruminants (for
instance, Bos moschatus) in the breeding season, many deer,
antelopes, sheep, and goats, possess odoriferous glands in various
situations, more especially on their faces. The so-called tear-sacs, or
suborbital pits, come under this head. These glands secrete a
semifluid fetid matter which is sometimes so copious as to stain the
whole face, as I have myself seen in an antelope. They are ‘usually
larger in the male than in the female, and their development is
checked by castration.’ According to Desmarest they are altogether
absent in the female of Antilope subgutturosa. Hence, there can be
no doubt that they stand in close relation with the reproductive
functions. They are also sometimes present, and sometimes absent,
in nearly allied forms. In the adult male musk-deer (Moschus
moschiferus), a naked space round the tail is bedewed with an
odoriferous fluid, whilst in the adult female and in the male until two
years old, this space is covered with hair, and is not odoriferous.”
Darwin believes in these cases that the odor serves to attract the
females. He admits that here, “active and long-continued use cannot
have come into play as in the case of the vocal organs.” He
concludes, therefore, that “the odor emitted must be of considerable
importance to the male, inasmuch as large and complex glands,
furnished with muscles for everting the sac, and for closing or
opening the orifice, have in some cases been developed. The
development of these organs is intelligible through sexual selection,
if the most odoriferous males are the most successful in winning the
females, and in leaving offspring to inherit their gradually perfected
glands and colors.”
There is sometimes a difference in the mammals in the hair of the
two sexes both in amount and in color. In some species of goats the
males have a beard, in others it is present in both sexes. The bull,
but not the cow, has curly hair on the forehead. In some monkeys
the beard is confined to the male, as in the orang; in other species it
is only larger in the males.
“The males of various members of the ox family (Bovidæ), and of
certain antelopes, are furnished with a dewlap, or great fold of skin
on the neck, which is much less developed in the female.
“Now, what must we conclude with respect to such sexual
differences as these? No one will pretend that the beards of certain
male goats, or the dewlap of the bull, or the crests of hair along the
backs of certain male antelopes, are of any use to them in their
ordinary habits.
“Must we attribute all these appendages of hair or skin to mere
purposeless variability in the male? It cannot be denied that this is
possible; for in many domesticated quadrupeds, certain characters,
apparently not derived through reversion from any wild parent form,
are confined to the males, or are more developed in them than in
the females—for instance, the hump on the male zebu cattle of
India, the tail of fat-tailed rams, the arched outline of the forehead
in the males of several breeds of sheep, and, lastly, the mane, the
long hairs on the hind-legs, and the dewlap of the male of the
Berbura goat.”
In these cases and in others that Darwin cites, which seem clearly
to indicate that some of these secondary sexual characters are not
the result of sexual selection, he concludes, “that they must be due
to simple variability, together with sexually limited inheritance.
 “Hence it appears reasonable to extend this same view to all
analogous cases with animals in a state of nature. Nevertheless I
cannot persuade myself that it generally holds good, as in the case
of the extraordinary development of hair on the throat and fore-legs
of the male Ammotragus, or in that of the immense beard of the
male Pithecia. Such study as I have been able to give to nature
makes me believe that parts or organs which are highly developed,
were acquired at some period for a special purpose. With those
antelopes in which the adult male is more strongly colored than the
female, and with those monkeys in which the hair on the face is
elegantly arranged and colored in a diversified manner, it seems
probable that the crests and tufts of hair were gained as ornaments;
and this I know is the opinion of some naturalists. If this be correct,
there can be little doubt that they were gained, or at least modified
through sexual selection; but how far the same view may be
extended to other mammals is doubtful.”
The astonishing colors in some of the monkeys cannot be passed
over without comment.
“In the beautiful Cercopithecus diana, the head of the adult male
is of an intense black, whilst that of the female is dark gray; in the
former the fur between the thighs is of an elegant fawn-color, in the
latter it is paler.
“In the Cercopithecus cynosurus and griseoviridis one part of the
body, which is confined to the male sex, is of the most brilliant blue
or green, and contrasts strikingly with the naked skin on the hinder
part of the body, which is vivid red.
“Lastly, in the baboon family, the adult male of Cynocephalus
hamadryas differs from the female not only by his immense mane,
but slightly in the color of the hair and of the naked callosities. In
the drill (C. leucophæus) the females and young are much paler-
colored, with less green, than the adult males. No other member in
the whole class of mammals is colored in so extraordinary a manner
as the adult male mandrill (C. mormon). The face at this age
becomes of a fine blue, with the ridge and tip of the nose of the
most brilliant red. According to some authors, the face is also
marked with whitish stripes, and is shaded in parts with black, but
the colors appear to be variable. On the forehead there is a crest of
hair, and on the chin a yellow beard. ‘Toutes les parties supérieures
de leurs cuisses et le grand espace nu de leurs fesses sont
également colorés du rouge le plus vif, avec un mélange de bleu qui
ne manque réellement pas d’élégance.’ When the animal is excited
all the naked parts become much more vividly tinted.”
Darwin sums up the evidence in regard to the differences in color
between the male and female in the following statement:—
“I have now given all the cases known to me of a difference in
color between the sexes of mammals. Some of these may be the
result of variations confined to one sex and transmitted to the same
sex, without any good being gained, and therefore without the aid of
selection. We have instances of this with our domesticated animals,
as in the males of certain cats being rusty-red, whilst the females
are tortoise-shell colored. Analogous cases occur in nature: Mr.
Bartlett has seen many black varieties of the jaguar, leopard, vulpine
phalanger, and wombat; and he is certain that all or nearly all these
animals, were males. On the other hand, with wolves, foxes, and
apparently American squirrels, both sexes are occasionally born
black. Hence it is quite possible that with some mammals a
difference in color between the sexes, especially when this is
congenital, may simply be the result, without the aid of selection, of
the occurrence of one or more variations, which from the first were
sexually limited in their transmission. Nevertheless it is improbable
that the diversified, vivid, and contrasted colors of certain
quadrupeds, for instance, of the above monkeys and antelopes, can
thus be accounted for.”
Finally, the case of man must be considered from the point of view
of sexual selection, for Darwin claims that man has acquired a
number of his secondary sexual characters in this way. For instance,
the beard is an excellent case of a secondary sexual character.
Darwin’s interpretation is that the beard has been retained, or even
developed, through the selection by the females of those males that
had this outgrowth best developed. Conversely, the absence of hair
on the face of the female is supposed by Darwin to have been
brought about by men selecting those women having less hair on
their faces. The greater intellect, energy, courage, pugnacity, and
size of man are the outcome of the competition of the males with
each other, since the individual excelling in these qualities will be
able to select the most desirable wife, or wives, and it is assumed
will, therefore, leave more descendants. The standard of beauty has
been kept up by men selecting the most beautiful women in each
generation (the fate of the other married women is ignored), and
this beauty is supposed to have been transmitted primarily to their
daughters, but also to their sons.
Although all these forms of selection are imagined to be acting in
man, either alternately or simultaneously, yet Darwin recognizes in
man a number of checks to the action of sexual selection: amongst
savages, the so-called communal marriages; second, infanticide,
generally of the young females, which appears in some races to be
practised to an astonishing degree; third, early betrothals; fourth,
the holding of women as slaves.
When we recall that selection to be effective can only be carried
out under very exacting conditions, we cannot but be appalled at the
demands made here on our credulity. The choice of the women has
produced the beard of man, the choice of man the absence of a
beard in women; the competition of the males with each other is
leading at the same time to the development of at least half a dozen
qualities that are supposed to be male specialities, and while all this
is going on the results are being checked sometimes by one means,
sometimes by another. Moreover, even this is not all that we are
asked to accept, for there are several other qualities of the male that
are put down as secondary sexual characters. For example, let us
examine what Darwin has to say in regard to the development of the
voice, and of singing in man.
 In man the vocal cords are about a third longer than in woman
and his voice deeper. Emasculation arrests the development of the
vocal apparatus, and the voice remains like that of a woman. This
difference between the sexes, Darwin thinks, is due probably to
long-continued use by the male “under the excitement of love, rage,
and jealousy.” In other words, an appeal is again made to the
Lamarckian theory, and in this case to explain the origin of an organ
that conforms to all the requirements of the secondary sexual
characters.
“The capacity and love for singing, or music, though not a sexual
character in man,” in the sense of being confined to one sex, yet is
supposed to have arisen through sexual selection in the following
way: “Human song is generally admitted to be the basis or origin of
instrumental music. As neither the enjoyment nor the capacity of
producing musical notes are faculties of the least use to man in
reference to his daily habits of life, they must be ranked amongst the
most mysterious with which he is endowed.”
Man is supposed to have possessed this faculty of song from a
very remote time, and even the most savage races make musical
sounds, although we do not enjoy their music, or they ours.
“We see that the musical faculties, which are not wholly deficient
in any race, are capable of prompt and high development, for
Hottentots and Negroes have become excellent musicians, although
in their native countries they rarely practise anything that we should
consider music. Hence the capacity for high musical development,
which the savage races of man possess, may be due either to the
practice by our semi-human progenitors of some rude form of music,
or simply to their having acquired the proper vocal organs for a
different purpose. But in this latter case we must assume, as in the
above instance of parrots, and as seems to occur with many
animals, that they already possessed some sense of melody.”
Darwin sums up the evidence in the two following statements, the
insufficiency of which to explain the phenomena is I think only too
obvious: “All these facts in respect to music and impassioned speech
become intelligible to a certain extent, if we assume that musical
tones and rhythm were used by our half-human ancestors, during
the season of courtship, when animals of all kinds are excited not
only by love, but by the strong passions of jealousy, rivalry, and
triumph. From the deeply laid principle of inherited associations,
musical tones in this case would be likely to call up vaguely and
indefinitely the strong emotions of a long past age.” Thus the
difficulty is shifted to the shoulders of our long-lost savage
ancestors; or even, in fact, to our simian forefathers, as the
following paragraph indicates:—
“As the males of several quadrumanous animals have their vocal
organs much more developed than in the females, and as a gibbon,
one of the anthropomorphous apes, pours forth a whole octave of
musical notes and may be said to sing, it appears probable that the
progenitors of man, either the males or females or both sexes,
before acquiring the power of expressing their mutual love in
articulate language, endeavored to charm each other with musical
notes and rhythm. So little is known about the use of the voice by
the Quadrumana during the season of love, that we have no means
of judging whether the habit of singing was first acquired by our
male or female ancestors. Women are generally thought to possess
sweeter voices than men, and as far as this serves as any guide, we
may infer that they first acquired musical powers in order to attract
the other sex. But if so, this must have occurred long ago, before
our ancestors had become sufficiently human to treat and value their
women merely as useful slaves. The impassioned orator, bard, or
musician, when with his varied tones and cadences he excites the
strongest emotions in his hearers, little suspects that he uses the
same means by which his half-human ancestors long ago aroused
each other’s ardent passions during their courtship and rivalry.”
We have now examined in some detail the evidence that Darwin
has brought forward in support of his hypothesis of sexual selection.
A running comment has been made while considering the individual
cases, but it may be well to sum up the matter by briefly indicating
the reasons why the hypothesis seems incompetent to explain the
facts.
 General Criticism of the Theory of Sexual Selection
1. Some of the objections that apply to the theory of natural
selection apply also with equal force to the theory of sexual selection
in so far as the results in both cases are supposed to be the
outcome of the selection of individual, or fluctuating, variations. If
these variations appear in only a few individuals, their perpetuation
is not possible, since they will soon disappear through crossing. It
would be, of course, preposterous to suppose that at any one time
only those few individuals pair and leave descendants that have the
secondary sexual characters developed to the highest point, but if
something of this sort does not occur, the extreme of fluctuating
variations cannot be maintained. Even if half of the individuals are
selected in each generation, the accumulation of a variation in a
given direction could not go very far. The assumption, however, that
only half of all the individuals that reach maturity breed, and that all
of these are chosen on account of the special development of their
secondary sexual characters, seems preposterous. Furthermore, if it
is assumed that the high development of the new character appears
in a large number of individuals, then it is not improbable that its
continued appearance might be accounted for without bringing in, at
all, the hypothesis of sexual selection.
2. But even supposing that the females select the most beautiful
males, then, since in the vast majority of higher animals the males
and the females are in equal numbers, the others will also be able to
unite with each other in pairs after this first selection has taken
place. Nothing will therefore be gained in the next generation. It is
interesting to see how Darwin attempts to meet this argument. He
tries to show in the case of birds, that there are always unpaired
individuals, but since the few facts that he has been able to collect
show that there are as many additional females as males, the
argument proves too much. A few species are polygamous, one male
having a number of female birds; but on this basis we can only
account, at best, for the development through competition of the
organs of offence and defence used to keep away the weaker males.
Yet it is just amongst these birds that we often find the ornamental
characters well developed. In fact, since all the females in such
cases are selected, and since they will transmit the characters of all
the males, it is evident that the secondary sexual characters could
not be formed in the way imagined.
3. If the female fails to select only the more ornamental males, no
result will follow. It has not been shown that she is capable of
making such a choice, and in the lower forms particularly, it does not
seem probable that this is done. The argument that Darwin often
employs, namely, that unless she does select, the display of the
males before her is meaningless, is not to the point. So far as we
can detect the “cause” of the display of the male, it appears to be
due to his own excitement; and even if we go so far as to admit that
the “purpose” is to attract the other sex, it still does not in the least
follow that the most ornamental male is selected, and unless this
occurs the display has no bearing on the hypothesis of sexual
selection.
4. The two forms of sexual selection, namely, competition of the
males with one another (really one form of natural selection), and
the selection of the most ornamental or gifted individuals, are both
used by Darwin to explain secondary sexual characters, the one for
organs of offence and defence, and the other for ornamental
characters. If we fully appreciate the difficulties that any theory of
selection meets with, we shall realize how extraordinarily complex
the action must be, when two such processes are carried out at the
same time, or even during alternating periods.
5. It has been objected to Darwin’s theory of sexual selection, that
he suddenly reverses its mode of action to explain those cases in
which the female is the stronger and more ornamented sex; but if,
as Darwin shows, the instincts of the male have also changed, and
have become more like those of the female, I can see no inherent
difficulty in this way of applying the theory. A much more serious
objection, it seems to me, is that the male is supposed to select the
female for one set of characteristics, and the female to select the
male for another set. It sounds a little strange to suppose that
women have caused the beard of man to develop by selecting the
best-bearded individuals, and the compliment has been returned by
the males selecting the females that have the least amount of beard.
It is also assumed that the results of the selection are transmitted to
one sex only. Unless, in fact, the character in question were from the
beginning peculiar to only one sex as to its inheritance, the two
sexes might go on forever selecting at cross-purposes, and the result
would be nothing.
6. The development, or the presence, of the æsthetic feeling in
the selecting sex is not accounted for on the theory. There is just as
much need to explain why the females are gifted with an
appreciation of the beautiful, as that the beautiful colors develop in
the males. Shall we assume that still another process of selection is
going on, as a result of which those females are selected by the
males that appreciate their unusual beauty, or that those females
whose taste has soared a little higher than that of the average (a
variation of this sort having appeared) select males to correspond,
and thus the two continue heaping up the ornaments on one side
and the appreciation of these ornaments on the other? No doubt an
interesting fiction could be built up along these lines, but would any
one believe it, and, if he did, could he prove it?
Darwin assumes that the appreciation on the part of the female is
always present, and he thus simplifies, in appearance, the problem,
but he leaves half of it unexplained.
7. It has been pointed out, that it is important to distinguish
between the possible excitement of the female by the display or
antics of the male, and the selection of the more beautiful or agile
performer. Darwin himself records a few cases, which plainly show
that the more beautiful is not always the more successful. It has also
been suggested that the battles of the males are sometimes sham
performances, and even when they are real, if the less vigorous do
not remain to be destroyed but run away, they live to find mates of
their own. In fact, the conduct of the males at the breeding season
appears to be much more the outcome of their own excitement than
an attempt to attract the females.
8. There is another side to the question, the importance of which
is so great, that it is surprising that Darwin has not taken any notice
of it. If, in order to bring about, or even maintain, the results of
sexual selection, such a tremendous elimination of individuals must
take place, it is surprising that natural selection would not counteract
this by destroying those species in which a process, so useless for
the welfare of the species, is going on. It is curious that this has not
been realized by those who believe in both of these two hypotheses.
9. What has just been said applies also with almost equal force to
the development of such structures as the horns of deer, bison,
antelopes, and the brilliant colors of many insects and birds. If in
nature, competition between species takes place on the scale that
the Darwinian theory of natural selection postulates, such forms, if
they are much exposed, would be needlessly reduced in numbers in
the process of acquiring these structures. So many individuals would
have been at such a disadvantage in breeding, that if competition is
as severe as the theory of natural selection postulates, these species
could hardly be expected to compete successfully with other species
in which sexual selection was not taking place.
10. Darwin admits that, in certain cases, external conditions may
have acted directly to produce the colors in certain forms, and if
these were not injurious he thinks they might have become
constant. Such cases are left unexplained in the sense that they are
not supposed to be adaptations to anything in particular. That colors
produced in this way might afterward be found useful, irrespective of
how they arose, is admitted as one of the ways in which sexual
differences may have arisen.
11. It is baffling to find Darwin resorting to the Lamarckian
explanation in those cases in which the improbability of the
hypothesis of sexual selection is manifest. If either principle is true,
we should expect it to apply to all phenomena of the same sort; yet
Darwin makes use of the Lamarckian principle, in the hypothesis of
sexual selection, only when difficulties arise.
12. In attempting to explain the development of the musical sense
in man, it is clear that the hypothesis of sexual selection fails to give
a satisfactory explanation. To suppose that the genius of a
Beethoven or of a Mozart could have been the result of a process of
sexual selection is too absurd to discuss. Neither the power of
appreciation nor of expression in music could possibly have been the
outcome of such a process, and it does not materially help the
problem to refer it back to a troop of monkeys making the woods
hideous with their cries.
We come now to some of the special cases to which Darwin’s
hypothesis has been applied.
13. In one case at least, it is stated that a bird living on the
ground might have acquired the color of the upper surface of the
body through natural selection, while the under surface of the males
of the same species might have become ornamented through the
action of sexual selection. Thus in one and the same individual the
two processes are supposed to have been at work, and it does not
lessen the difficulty very much by supposing the two processes to
have been carried out at different times, because it is evident that
what had been gained at one time by one process might become
lost while the color of certain parts was being acquired through the
other process.
14. Darwin points out that “the plumage of certain birds goes on
increasing in beauty during many years after they are fully mature,”
as in the peacock, and in some of the birds of paradise, and with the
plumes and crests of some herons. This is explained as possibly
merely the result of “continued growth.” The improbability of
selection is manifest in these cases, but if “continued growth” can
accomplish this much, why may not the whole process be also the
outcome of such growth? At any rate, whatever the explanation is, it
is important to find a case of a secondary sexual character that the
hypothesis obviously is insufficient to explain.
15. It is admitted in a number of cases, as in the stag for instance,
that, although the larynx of the male is enlarged, this is not, in all
probability, the outcome of sexual selection, but in other forms this
same enlargement is ascribed to the selection process.
16. It is admitted that in none of the highly colored British moths
is there much difference according to sex, although when a
difference of color is found in butterflies this is put down to the
action of sexual selection. If such wonderful colors as those of moths
can arise without the action of selection, why make a special
explanation for those cases in which this difference is associated
with sex?
17. It is well known that birds sing at other times of the year than
at the breeding season, and an attempt is made to account for this
in that birds take pleasure in practising those instincts that they
make use of at other times, as the cat plays with the captive mouse.
Does not this suggest that, if they had certain instincts, they would
be more likely to employ them at the times when their vitality or
excitement is at its highest without regard to the way in which they
have come by them?
18. The color of the iris of the eyes of many species of hornbills is
said to be an intense crimson in the males, and white in the females.
In the male condor the eye is yellowish brown, and in the female a
bright red. Darwin admits that it is doubtful if this difference is the
result of sexual selection, since in the latter case the lining of the
mouth is black in the males, and flesh-colored in the females, which
does not affect the external beauty. Yet if these colors were more
extensive and on the exterior, there can be little doubt that they
would have been explained as due to sexual selection.
19. When the females in certain species of birds differ more from
each other than they do from their respective males, the case is
compared to “those inexplicable ones, which occur independently of
man’s selection in certain sub-breeds of the game-fowl, in which the
females are very different, whilst the males can hardly be
distinguished.” Here then is a case of difference in color associated
with sex, but not the outcome of sexual selection.
20. The long hairs on the throat of the stag are said possibly to be
of use to him when hunted, since the dogs generally seize him by
the throat, “but it is not probable that the hairs were specially
developed for this purpose; otherwise the young and the females
would have been equally protected.” Here also is a sexual difference
that can scarcely be ascribed to selection.
Some cases of differences in color between the sexes “may be the
result of variations confined to one sex, and transmitted to the same
sex without any good being gained, and, therefore, without the aid
of selection. We have instances of this with our domesticated
animals, as in the males of certain cats being rusty-red while the
females are tortoise-shell colored. Analogous cases occur in nature:
Mr. Bartlett has seen many black varieties of the jaguar, leopard,
vulpine phalanger, and wombat; and he is certain that all or nearly
all of these animals were males.” If changes of this sort occur,
associated with one sex, why is there any need of a special
explanation in other cases of difference?

In the light of the many difficulties that the theory of sexual

selection meets with, I think we shall be justified in rejecting it as an
explanation of the secondary sexual differences amongst animals.
Other attempts to explain these differences have been equally
unsuccessful. Thus Wallace accounts for them as due to the
excessive vigor of the male, but Darwin’s reply to Wallace appears to
show that this is not the cause of the difference. He points out that,
while the hypothesis might appear plausible in the case of color, it is
not so evident in the case of other secondary sexual characters,
such, for instance, as the musical apparatus of the males of certain
insects, and the difference in the size of the larynx of certain birds
and mammals.
 Darwin’s theory served to draw attention to a large number of
most interesting differences between the sexes, and, even if it prove
to be a fiction, it has done much good in bringing before us an array
of important facts in regard to differences in secondary sexual
characters. More than this I do not believe it has done. The theory
meets with fatal objections at every turn.
In a later chapter the question will be more fully discussed as to
the sense in which these secondary sexual differences may be
looked upon as adaptations.
 CHAPTER VII

THE INHERITANCE OF ACQUIRED CHARACTERS AS A FACTOR

IN EVOLUTION

Lamarck’s Theory

One of the most striking and peculiar characteristics of living things

is that through use a part is able to carry out a particular function
better than before, and in some cases the use of the part leads to its
increase in size. Conversely, disuse leads to the decrease of a part in
size. We are perfectly familiar with this process in ourselves as
applied to our nervous system and muscles.
It is not surprising that the idea should have arisen that, if the
results of the use of a part are inherited by the next generation, the
adaptation of organisms might be explained in this way. The
presence of the organs of touch, in those parts of the body that are
more likely to come into contact with foreign bodies, offers a striking
parallel to the perfecting of the sensation of touch that can be
brought about through the use of any part. The development of eyes
only on the exposed parts of the body, as on the tentacles of the
sedentary annelids, or along the margin of the mantle of a bivalve
mollusk, suggests that there may be some direct connection
between their presence in these regions and the effect of light on
the parts. In fact, ever since the time of Lamarck, there have been
many zoologists who have claimed that many of the adaptations of
organisms have arisen in this way, that is, through the inheritance of
the characters acquired through use. In general this theory is
summed up in the phrase, “the inheritance of acquired characters.”
This view is prominently associated with the name of Lamarck,
who held, however, a different view in regard to the origin of some
of the other structures of the organism. Moreover, Erasmus Darwin,
even before Lamarck, had suggested the principle of the inheritance
of acquired characters.
As has just been said, Lamarck held that the inheritance of
acquired characters was only one of the ways in which animals have
become changed, and he clearly stated that in the case of all plants
and of some of the lower animals the change (evolution) which he
supposed them to undergo was due to the general influence of the
environment. Since plants and the lower animals (as he supposed)
have no central nervous system, or at least no such well-defined
nervous system as have the higher animals, Lamarck thought that
they could not have evolved in the same way as have the higher
animals. We now know that, so far as the lower animals, at least,
are concerned, there was no need for such a distinction, since many
of their responses are like those of the higher animals. This
distinction that Lamarck made is responsible, no doubt, for a
misconception that was long held in regard to a part of his views. It
is often stated that he supposed the desire of the animal for a
particular part has led to the development of that part; while in
reality he only maintained the desire to use a particular organ to
fulfil some want led to its better development through exercise, and
the result was inherited. Lamarck also supposed that the decrease in
use of a part which leads to its decrease in size accounts for the
degeneration of organs.
Lamarck first advanced his theory in 1801, when he cited the
following examples in its favor. A bird, driven through want to the
water to find its food, will separate its toes when they strike the
water. The skin uniting the bases of the toes will be stretched in
consequence, and in this way the broad membrane between the
toes of ducks and geese has been acquired. The toes of a bird that
is in the habit of perching on a tree become elongated in
consequence of becoming stretched, hence has arisen the foot with
the long toes characteristic of arboreal birds.
Shore-birds, “which do not care to swim,” but must approach the
water in order to obtain food, will be in danger of sinking into the
mud, “but, wishing to act so that their body shall not fall into the
liquid, they will contract the habit of extending and lengthening their
legs.” Hence have arisen the stiltlike legs of shore-birds.
These ideas were more fully elaborated in the following year. He
added the further examples: Our dray-horses have arisen through
the use to which they have been put, and the race-horse also, which
has been used in a different way. Cultivated plants, on the contrary,
are the result of the new environment to which they have been
subjected.
In the “Philosophic Zoologique,” published in 1809, Lamarck has
much more fully developed his theory. Here he combats strenuously
the idea that species are fixed. His point of view may be judged by
the following propositions, which he believes can be established:—
1. That all organized bodies of our globe are veritable productions
of nature, which she has successively produced in the course of a
long time.
2. That in her progress nature began, and begins still every day, to
produce the simplest organisms, and that she still produces directly
the same primitive kinds of organizations. This process has been
called spontaneous generation.
3. That the first beginning of animals and of plants takes place in
favorable localities and under favorable circumstances. An organic
movement having once established their production, they have of
necessity gradually developed their organs, and have become
diversified in the course of time.
4. That the power of growth of each part of the body being
inherited as a consequence of the first effect of life, different modes
of multiplication and of regeneration have arisen, and these have
been conserved.
5. That with the aid of sufficient time and of favorable
circumstances the changes that have taken place on the surface of
the globe have called forth new structures and new habits, and in
consequence have modified the organs of the body, and made
animals and plants such as we see them at the present day.
6. Finally, as a result of these changes that living bodies have
been forced to undergo, species have been formed, but these
species have only a relative constancy, and are not as ancient as is
nature herself. If the environment remains the same, species also
remain the same, as is exemplified by the animals living at present
in Egypt, which are exactly like those living there in ancient times.
Lamarck concludes that the appearance of stability is always
mistaken by the layman for the reality, because, in general, every
one judges things relatively to himself. In fact, species are not
absolutely constant, but are so only temporarily. “The influence of
the environment is continuous and always active, but its effects may
only be recognized after a long time.” The irregularity and the
complexity of the organization of animals is the outcome of the
infinitely diversified circumstances to which they have been
subjected. These changes, Lamarck claims, do not directly cause
modifications in the form of animals,[17] but bring about changes in
their needs, and changes in their needs bring about changes in their
actions. If the needs remain the same, the acquired actions become
habits. These habitual actions lead to the use of certain parts in
preference to others, and this in turn to an alteration in form and
structure. The individuals so changed breed together and leave
descendants that inherit the acquired modification.
17. This is clearly meant to be applied only in the case of higher animals.
Curiously enough, Lamarck follows up this argument by citing
some cases amongst plants that have been changed directly by the
action of the environment. He says that since plants have no
motions they have consequently no habits, but they are developed
by changes in their nutrition, etc., and this brings about the
superiority of some of the vital movements over others.
Amongst domestic animals Lamarck cites the case of the dog, that
has come from a wild form like the wolf, but having been carried into
different countries has acquired different and new habits, and this
has led to the formation of new races, such as the bulldog,
greyhound, pug-dog, spaniel, etc.
Lamarck’s argument shifts so often back and forth from animals to
plants, that it is clear that in his own mind he did not see any
important difference between the action of the environment on
plants, and the use of the organs of the animal. He gives in this
same connection his oft-quoted summary of what he calls the two
laws of nature “which observation always establishes.”
First Law. In every animal, that has not passed beyond the term of
its development, the frequent and sustained use of any organ
strengthens it, develops it, increases its size, and gives it strength
proportionate to the length of time of its employment. On the other
hand, the continued lack of use of the same organ sensibly weakens
it; it deteriorates, and its faculties diminish progressively until at last
it disappears.
Second Law. Nature preserves everything that she has caused the
individual to acquire or to lose by the influence of the circumstances
to which the race has been for a long time exposed, and
consequently by the influence of the predominant use of certain
organs (or in consequence of its continued disuse). She does this by
the generation of new individuals which are produced with the newly
acquired organs. This occurs, provided that the acquired changes
were common to the two sexes, or to the individuals that produced
the new forms.
These laws are, Lamarck says, fundamental truths which cannot
be misunderstood except by those who have never observed or
followed nature in her operations. He insists that it is a mistake to
suppose that the parts are responsible for the functions, for it is easy
to demonstrate that it is the needs and uses of the organs that have
caused the parts to develop.
If it is supposed, he continues, that these laws are hypothetical,
they may be demonstrated by the following facts: The adult baleen
whale is without teeth, although in the fœtus teeth are present,
concealed in the jaws. The loss of the teeth is the result of the
whale swallowing its food without first masticating it. The ant-eater
is also without teeth, and has also the habit of swallowing its food
without chewing it. The mole has very small eyes, and this is the
result of its having made very little use of them, since its habits are
subterranean. Another animal, the aspalax, has only the rudiments
of eyes, and has almost completely lost the power of sight. This
animal also lives underground like the mole.
Proteus, an aquatic salamander living in deep caves, has only
rudimentary eyes. In these latter cases it is the disuse of the eye
that has led to its degeneration. This is proven, Lamarck adds, by
the fact that the organs of hearing are never in this condition,
because sound vibrations penetrate everywhere, even into the
densest bodies.
It is a part of the plan of organization of the reptiles that they
have four legs; but the snakes, although belonging to this group,
have no legs. This absence of legs is explained by their having
acquired the habit of gliding over the ground, and of concealing
themselves in the grass. Owing to their repeated effort to elongate
themselves, in order to pass through narrow spaces, their bodies
have become drawn out. Under these circumstances legs would be
useless, since long ones would interfere with their motion, and short
ones could not move their long bodies. Since the plan of
organization limits the snakes to only four legs, and since this
number would be useless, they have disappeared.
Many insects are destitute of wings, although wings are a part of
the plan of organization of this group. They are absent only in those
forms whose habits render wings useless, consequently they have
disappeared through disuse.
The preceding cases are those in which the disuse of an organ has
led to its degeneration. The following cases are cited to show that by
use an organ increases in size. The formation of the web in the feet
of water-birds has already been given as a character which Lamarck
supposes to have been acquired through use; also the case of shore-
birds, which, by an effort to elongate their legs, have actually made
them so in the course of time. The necks of water-birds are also long
on account of their having been stretched in the efforts to catch fish.
The long tongues of the ant-eater, of the woodpecker, and of
humming-birds are the result of use, and the long, forked tongue of
serpents has come from their using their tongue to feel objects in
front of them.
Fishes that have acquired the habit of living in shallow water,
flounders, soles, etc., have been forced to swim on their sides in
order to approach nearer to the shore. Since more light comes from
above than from below, the eye on the under side, straining to turn
to the light, has finally migrated to the upper side.
The habit of eating great quantities of food, which distends the
digestive organs, has caused the bodies of herbivorous quadrupeds
to become large, as seen in the elephant, the rhinoceros, oxen,
horses, and buffaloes. The habit of standing for a long time on their
feet has caused some animals to develop hard, thick hoofs.
Herbivorous animals, that inhabit countries where they are
constantly subjected to attack, as deer and antelopes for example,
are forced to escape by rapid flight, and in consequence their bodies
have become slenderer and their legs thinner. The horns, antlers,
and protuberances that many of these animals possess are the
results of their butting each other when angered.
“The long neck and the form of the giraffe offer a curious case.
We know that the giraffe is the tallest of all animals. It inhabits the
centre of Africa, living in those localities where the earth is nearly
always dry and without herbage. It is obliged to browse on the
foliage of trees, and this leads to its stretching continually upwards.
As a result of this habit, carried on for a long time, in all the
individuals of the race, the anterior limbs have become longer than
the posterior, and its neck has also lengthened, so that the giraffe
without rising on its hind-legs stretches up its neck and can reach to
the height of six metres.”
The curved claws of the carnivora have arisen from the necessity
of grasping their prey. The power of retracting the claws has also
been acquired by the effort to draw them in when running over hard
ground. The abdominal pouch of the kangaroo, in which the young
are carried, opens anteriorly, and this has led to the animal standing
erect so that its young are not injured. In consequence, the fore-legs
have become shorter through disuse, and the hind-legs have
become stronger through use. The tail, which is also used as a
support, has become enormously thick at its base.
The sloth has been compelled to seek refuge in the trees, and has
taken up its abode permanently there, feeding on leaves. Its
movements are limited to those involved in crawling along the limbs
in order to reach the leaves. After feeding it remains inactive and
sluggish, these habits being provoked by the heat of the climate.
The results of its mode of life have been to cause the arms to
become elongated due to the habit of the sloth of grasping the limbs
of the tree; the claws of the fingers and toes have also become long
and hooked in order to retain their hold. The digits that do not make
any individual movements have lost the power to do so, and have
become fused, and can only be bent in and straightened out. The
thighs, being bent out to clasp the larger branches, have caused the
pelvis to widen, and, in consequence, the cotyloid cavities have
become directed backward. Many of the bones of the skeleton have
become fused, as a result of the immobility of the animal.
Lamarck says, that “Nature, in producing, successively, all the
species of animals, beginning with the most imperfect, or the most
simple, and terminating with the most perfect, has gradually
complicated their organization. These animals becoming scattered
throughout the habitable regions of the globe each species has
received from the influences of its surroundings its present habits,
and the modifications of the parts the use of which we recognize.”
Such are Lamarck’s views and a fairly complete statement of the
facts from which he draws his conclusions. His illustrations appear
naïve, and often not a little ludicrous, but it must be admitted that,
despite their absurdities, his theory appears in some cases to
account wonderfully well for the facts. The long legs of wading birds,
the long neck and disproportionately long fore-legs of the giraffe,
the structure of the sloth, and particularly the degeneration of the
eyes of animals living in the dark, seem to find a simple explanation
in the principle of the inheritance of acquired characters. But the
crucial point of the entire theory is passed over in silence, or rather
is taken for granted by Lamarck, namely, the inheritance in the
offspring of the characters acquired through use or disuse in the
parent. He does not even discuss this topic, but in several places
states unreservedly that the increase or decrease of a part reappears
in the next generation. It is here that Lamarck’s theory has been
attacked in more modern times, for as soon as experimental proof
was demanded to show that the results of use or of disuse of an
organ is inherited, no such proof was forthcoming. Yet the theory is
one that has the great merit of being capable of experimental test,
and it is astonishing to find that, with the immense amount that has
been written by his followers, so few attempts have been made to
give the theory a thorough test. The few results that have been
obtained are not, however, favorable to the theory, but almost the
only attempts at experiment that have been made in this direction
have been those of mutilating certain parts; and were it not for
popular belief to the effect that such mutilations are inherited, one
would least expect to get evidence for or against the theory in this
direction. Lamarck himself believed that the changes were slowly
acquired, and I think modern Lamarckians are justified in claiming
that the validity of the theory can only be tested by experiments in
which the organism is subjected to influences extending over a
considerable period, although Lamarck appears to have believed that
the first results may appear quite soon. Before expressing any
opinion in regard to the probability of the theory, let us examine
what the followers of Lamarck have contributed in the way of
evidence to the theory, rather than the applications that they have
made of the theory. We shall also find it profitable to consider some
of the modern criticism, to which the theory has been subjected.
Despite the contempt with which Darwin referred to Lamarck’s
theory, he himself, as we have seen, often made use of the principle
of the inheritance of acquired characters, and even employed the
same illustrations cited by Lamarck. Darwin seems to have
misunderstood Lamarck’s view, and to have accepted the current
opinion that Lamarck supposed an animal acquired a new organ by
desiring or needing it. Darwin says, “Heaven forefend me from
Lamarck’s nonsense of a tendency to progressive adaptation from
the slow willing of the animals.” Darwin speaks of Lamarck as stating
that animals will that the egg shall be a particular form so as to
become attached to particular objects. Lamarck’s latest biographer,
Packard, says he is unable to find any statements of this sort in
Lamarck’s writings.
The following cases that Darwin tried to explain through the
inheritance of acquired characters are exactly like those to which
Lamarck applied his theory. The bones of the wing of the domestic
duck weigh less than those of the wild duck, and the bones of the
leg more. Darwin believes this is due to the effects of the inheritance
of acquired characters. The drooping ears of many domestic
mammals are also explained by him as a result of disuse—“the
animals being seldom much alarmed.” In speaking of the male of the
beetle, Onites apelles, Darwin quotes Kirby to the effect that the
tarsi are so habitually lost that the species has been described
without this part of the foot. In the sacred beetle of Egypt the tarsus
is totally absent. Hence he concludes that the absence of tarsi in the
sacred beetle, and the rudimentary condition of the tarsus in others,
is probably the result of disuse, rather than a case of inheritance of
a mutilation. Darwin grants that “the evidence that accidental
mutilations can be inherited is at present not decisive, but the
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