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 11create™
Course Global Business Today
Course Number Third Asia Pacific Edition
Hill
Cronk
Wickramasekera
McGraw-Hill Education
Copyrigllt © 2014 Mc6raw·Hill Education (Australia) Ply Ltd
Additional owners of copyright are ackn~ged in on·page credits
Every effort has been made to trace and acknowledge copyrighted material. The authors
and publishers tender their apologies should any infringement have occurred.
Reproduction and communication for educational purposes
The Australian Copyright Ad 1968 (the Act) allows a maximum of one chapter°' 10% of
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the attention of the permissions editor at the address below.

National Library of Australia Cataloguing·in·Publication Data

Author: Hill, Charles W. L, author.
Title: Global business today I Charles W.L Hill; Thomas Cronk;
Rumintha Wackramasekera.
Edttioo: 3rd edition.
ISBN: 9781743070154 (paperback)
Notes: Previous ed.: 2010.
Includes b ibliographical references and index.
Subjects: International business enterprises-Management
International trade .
Investments, Foreign.
Capital market
Olhe r Authors/
Contributors: Cronk, Thomas, author.
Wickramasekera, Rumintha, author.
Dewey Number: 658.049

Published in Australia by
McGraw-Hill Education (Australia) Ply Ltd
Level 2, 82 Waterloo Road, North Ryde NSW 2113
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For a list of countries and their capitals, see pages 6 ~.

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CONTENTS IN FULL
Preface xiv Text at a Glance xx
About the Authors xv Case and Vignette Matrix xxii
Acknowledgments xvi Digital Resources xxvi
What's New in the Third Edition xvii International Business Graduate Attributes xxviii
Organisation/Content xviii

PART ONE GLOBALISATION 1

CHAPTER 1 GLOBALISATION 2
OPENING CASE: KOMATSU: GLOBALISATION The changing wortd order 41
THROUGH DECISIVELY BEITER BUSINESS 2 The global economy of the 21st century:
International Business Graduate Attributes (IBGA) 6 the emerging markets century? 43
Learning Objectives 7 The Globalisation Debate 48
Introduction 7 Anti-globalisation protests 48
What is Globalisation? 12 Globalisation, jobs and income inequality 49
The globalisation of markets 12 Globalisation, labour policies and the environment 53
The globalisation of production 13 Globalisation and national sovereignty 57
The Emergence of Global Institutions 17 Globalisation and the world's poor 59
Drivers of Globalisation 19 Managing in the Global Marketplace:
Declining trade and investment barriers 19 What's the Difference? 61
The role of technological change 23 Key Terms 63
Implications for the globalisation of production Summary 63
and markets 26 International Business Graduate Attributes:
The Changing Shape of the Global Economy 28 Learning and Assessment Tasks 65
The changing world output and world trade picture 29 CLOSING CASE: SHOES OF PREY 66
The changing foreign direct investment picture 32 Endnotes 68
The changing nature of the multinational enterprise 37

PART TWO CROSS-BORDER LINKAGES:

TRADE, INVESTMENT AND EXCHANGE 73
CHAPTER 2 THEORIES OF TRADE, INVESTMENT
AND INTERNATIONALISATION 74
OPENING CASE: BANGLADESH'S TEXTILE TRADE 74 Heckscher-Ohlin theory 86
International Business Graduate Attributes (IBGA) 75 The Leontief paradox 87
Learning Objectives 76 The Product Life-Cycle Theory 88
Introduction 76 Evaluating the product life-cycle theory 88
Mercantil ism 77 New Trade Theory 90
Absolute Advantage 78 Increasing product variety and reducing costs 90
Comparative Advantage 79 Economies of scale, first-mover advantages
Ricardo's theory 79 and the pattern of trade 91
The Samuelson critique 84 Implications of new trade theory 92
Evidence for the link between trade and growth 84 National Competitive Advantage: Porter's Diamond 94
viii I CONTENTS I
Factor endowments 94 The Uppsala Models (LI-Models) 103
Demand conditions 95 The Innovation Models (I-Models) 104
Related and supporting industries 95 International New Ventures/Born Global Firms 105
Firm strategy, structure and rivalry 96 Focus on Managerial Implications 106
Evaluating Porter's theory 96 Key Terms 108
Foreign Direct Investment in the World Economy 98 Summary 108
Why foreign direct investment? 98 International Business Graduate Attributes:
The pattern of foreign direct investment 100 Learning and Assessment Tasks 109
The eclectic paradigm 101 CLOSING CASE: LOG ITECH 111
The Stage Models of Internationalisation: Endnotes 112
An Incremental Sequential Approach 103

CHAPTER 3 THE POLITICAL ECONOMY OF

TRADEANDINVESTMENT 116
OPENING CASE: UNEASE OVER CHINA'S RARE Government Policy Instruments and FOi 140
EARTHS TRADE POLICIES 116 Home-country policies 140
International Business Graduate Attributes (IBGA) 119 Host-country policies 141
Learning Objectives 119 Trade and FOi liberalisation 145
Introduction 119 Trade and FOi liberalisation and the WTO 148
Instruments of Trade Policy 121 Regional Economic Integration 157
Tariffs 121 The move towards regional economic integration 157
Subsidies 123 Levels of economic integration 159
Import quotas and voluntary export restraints 124 The case for regional economic integration 161
Local content requirements 125 The case against regional economic integration 162
Administrative policies 126 Focus on Managerial Implications 163
Antidumping policies 126 Key Terms 165
Why Governments Intervene in Trade 127 Summary 166
Political arguments for intervention 127 International Business Graduate Attributes:
Economic arguments for intervention 131 Learning and Assessment Tasks 167
Why Governments Intervene in FOi 136 CLOSING CASE: TOBACCO PLAIN-PACKAGING LAWS
Host-country benefits and costs 137 IN AUSTRALIA: HEALTH DR TRADE ISSUES? 169
Home-country benefrts and costs 139 Endnotes 170

CHAPTER 4 FOREIGN EXCHANGE AND THE

INTERNATIONAL MONETARY SYSTEM 174
OPENING CASE: BILLABDNG 174 Investor psychology and bandwagon effects 192
International Business Graduate Attributes (IBGA) 176 The International Monetary System 192
Learning Objectives 176 Exchange rate regimes in practice 192
Introduction 176 Motives and means for managing the foreign
The Foreign Exchange Market 178 exchange rate 197
The nature of the foreign exchange market 178 Functions of an international monetary system 199
The functions of the foreign exchange market 179 The rise and fall of the Breiten Woods system 200
Determination of the Exchange Rate 188 A role for the IMF 203
Prices and exchange rates 188 Focus on Managerial Implications 212
Interest rates and exchange rates 190 Key Terms 216
 I CONTENTS I ix

Summary 217 CLOSING CASE: THE EUROZONE CRISIS:

International Business Graduate Attributes (IBGA): UNFINISHED BUSINESS 220
Learning and Assessment Tasks 218 Endnotes 224

PART THREE COUNTRY DIFFERENCES 227

CHAPTER 5 DIFFERENCES IN CULTURE 228
OPENING CASE: PANASON IC ANO JAPAN'S Superstitions 249
CHANGING CULTURE 228 Language 251
International Business Graduate Attributes (IBGA) 230 Spoken language 251
Learning Objectives 230 Unspoken language 252
Introduction 231 Education 253
What is Culture? 231 Culture and the Workplace 254
Values and norms 232 Cultural context 254
The determinants of culture 234 Cultural Change 256
Social Structure 235 Focus on Managerial Implications 258
Individuals and groups 235 Key Terms 261
Social stratification 237 Summary 261
Religious and Ethical Systems 241 International Business Graduate Attributes (IBGA):
Christianity 241 Learning and Assessment Tasks 262
Islam 243 CLOSING CASE: COGNITION CONSULTING:
Hinduism 245 EXPORTING NEW ZEALAND'S EDUCATIONAL
Buddhism 247 EXPERTISE AND CULTURE? 263
Confucianism 248 Endnotes 265

CHAPTER 6 POLITICAL AND LEGAL ENVIRONMENTS 268

OPENING CASE: THE RISKS OF REGULATORY Differences in contract law 294
CHANGE: ANIMAL WELFARE AND THE LIVE Property rights and cmuption 295
CATILE TRADE TO INOONESIA 268 The protection of intellectual property 298
International Business Graduate Attributes (IBGA) 270 Product safety and product liability 300
Learning Objectives 271 Competition law 302
Introduction 271 Focus on Managerial Implications 305
Political Systems 272 Key Terms 308
Collectivism and individualism 273 Summary 309
Shifting ideology and FOi 278 International Business Graduate Attributes:
Democracy and totalitarianism 283 Learning and Assessment Tasks 310
The spread of democracy 285 CLOSING CASE: GROWING AUTHORITARIAN ISM
The new world order and global terrorism 289 AND BUSINESS IN FIJ I 311
Legal Systems 293 Endnotes 315
Different legal systems 293

CHAPTER 7 ECONOMIC ENVIRONMENT 318

OPENING CASE: POLI SH ECONOMIC RESILI ENCE 318 Learning Objectives 320
International Business Graduate Attributes (IBGA) 320 Introduction 320
• I CONTENTS I
Economic Endowments-Natural and Created 321 Market economy 346
Size 321 Command economy 346
Geography 321 Mixed economy 347
People 322 Economic systems and economic development 348
Infrastructure and institutions 326 Economies in Transition 353
Productivity and competiti~ness 327 The spread of market-based systems 353
Economic Performance-Macroeconomic Stability 329 The nature of economic transition 354
Economic stability 329 Focus on Managerial Implications 359
Economic growth and employment fluctuations 331 Key Terms 363
Inflation 333 Summary 363
External viability and the balance of payments 335 International Business Graduate Attributes(IBGA):
Economic Performance-Economic Development 338 Learning and Assessment Tasks 364
Measures of economic development 338 CLOSING CASE: EMBRAER FLI ES AS BRAZIL'S
Broader conceptions and measures of development 343 EXPORTS HIT HEADWINDS 366
Economic Systems 345 Endnotes 370

CHAPTER 8 ETHICS AND CORPORATE

RESPONSIBILITY 372
OPENING CASE: EXECUTIVES ACTING The Friedman doctrine 390
UNETHICALLY: COMPANIES PENALISED 372 Cultural relativism 391
International Business Graduate Attributes (IBGA) 374 Righteous moralism 392
Learning Objectives 374 Naive imm()(alism 393
Introduction 374 Utilitarian and Kantian ethics 393
Ethical Issues in International Business 375 Rights theories 394
Employment practices 375 Justice the()(ies 396
Human rights 376 Ethical Decision Making in International Business 397
Environmental pollution 378 Organisation culture and leadership 398
Bribery and cmuption 380 Decision-making processes 399
Moral obligations 382 Ethics officers 401
Ethical Dilemmas 383 Moral courage 401
Child labour 384 Summary of managerial actions 401
The Roots of Unethical Behaviour 385 Key Terms 402
Personal ethics 385 Summary 402
Decision-making processes 386 International Business Graduate Attributes (IBGA):
Organisation culture 386 Learning and Assessment Tasks 403
Unrealistic perfonmance expectations 387 CLOSING CASE: SIEMENS BRIBERY SCANDAL 404
Corporate go~rnance and leadership 388 Endnotes 405
Philosophical Approaches to Ethics 390

CHAPTER 9 COUNTRY MARKET ANALYSIS 408

OPENING CASE: VELLUS PET COSMETICS 408 Improving Exp()(! Performance 412
International Business Graduate Attributes (IBGA) 409 An international comparison 412
Learning Objectives 410 Export promotion agencies 413
Introduction 410 Utilising export management companies 414
The Promise and Pitfalls of Exporting 410 Export strategy 414
 CONTENTS I xi

Basic Entry Decisions 415 Bill of lading 429

Which foreign markets? 415 A typical international trade transaction 431
Gross fixed capital formation 420 Export credit insurance 432
The cage framework for distance analysis 421 Summary 432
Cost-risk trade-off 422 Key Terms 434
Timing of entry 423 Summary 435
Scale of entry and strategic commitments 424 International Business Graduate Attributes (IBGA):
Financing Trade 426 Leaming and Assessment Tasks 435
Lack of trust 427 CLOSING CASE: UNIQLO GOES GLOBAL 436
Letter of credit 428 Endnotes 438
Draft 429

PART FOUR COMPETING IN THE

GLOBAL MARKETPLACE 441
CHAPTER 10 THE STRATEGY OF
INTERNATIONAL BUSINESS 442
OPENING CASE: AVON PRODUCTS 442 Differences in distribution channels 466
International Business Graduate Attributes (IBGA) 444 Host government demands 467
Leaming Objectives 444 Choosing a Strategy 46 7
Introduction 444 Global standardisation strategy 469
Strategy and the Firm 445 Localisation strategy 470
Value creation 446 Transnational strategy 470
Strategic positioning 447 International strategy 471
Operations: the firm as a value chain 449 The Evolution of Strategy 472
Organisation: the implementation of strategy 452 Strategic Alliances 473
In sum: strategic fit 453 The advantages of strategic alliances 473
Global expansion, profitability and profit growth 454 The disadvantages of strategic alliances 475
Expanding the market: leveraging products Making alliances work 476
and competencies 454 Key Terms 4 79
Location economies 456 Summary 4 79
Experience effects 458 International Business Graduate Attributes (IBGA):
Leveraging subsidiary skills 460 Leaming and Assessment Tasks 480
Cost Pressures and Pressures for Local CLOSING CASE: THE GLOBALISATION OF A WINE
Responsiveness 463 BRAND FROM AN EMERGING MARKET: THE CASE
Pressures for cost reductions 464 OF CASILLERO DEL DIAB LO 481
Pressures for local responsiveness 465 Endnotes 483

CHAPTER 11 ENTERING FOREIGN MARKETS 486

OPEN ING CASE: GENERAL MOTORS' JOINT Exporting 489
VENTURE IN CH INA 486 Importing 492
International Business Graduate Attributes (IBGA) 487 Turnkey projecls 492
Leaming Objectives 488 Licensing 493
Introduction 488 Franchising 495
Entry Modes 488 Joinl ventures 496
xii I CONTENTS I
Wholly owned subsidiaries 499 Types of countertrade 508
Selecting an Entry Mode 500 The pros and cons of countertrade 509
Core competencies and entry mode 500 Key Terms 510
Pressures for cost reductions and entry mode 502 Summary 510
Greenfield Venture Versus Acquisition 502 International Business Graduate Attributes (IBGA):
Pros and cons of acquisitions 502 Learning and Assessment Tasks 511
Pros and cons of greenfield ventures 506 CLOSING CASE: TESCO ACH IEVES INTERNATIONAL
Making a choice: greenfield or acquisition? 506 SUCCESS 512
Countertrade 507 Endnotes 513
The incidence of countertrade 507

CHAPTER 12 INTERNATIONAL MARKETING

AND R&D 516
OPENING CASE: FORD'S GLOBAL MARKETING 516 Strategic pricing 538
International Business Graduate Attributes (IBGA) 517 Predatory pricing 538
Learning Objectives 518 Regulatory influences on prices 539
Introduction 518 Configuring the Marketing Mix 540
The Globalisation of Markets and Brands 519 New Product Development 541
Market Segmentation 521 The location of R&D 542
Product Attributes 522 Integrating R&O, marketing and p<0duction 543
Cultural differences 522 Cross-functional teams 544
Economic development 523 Building global R&O capabilities 545
Product and technical standards 523 Key Terms 546
Distribution Strategy 524 Summary 546
Differences between countries 524 International Business Graduate Attributes:
Choosing a distribution strategy 527 Learning and Assessment Tasks 547
Communication Strategy 528 CLOSING CASE: RISKY BUSINESS:
Barriers to international communication 528 AN AUSTRALIAN FIRM IN CONFLICT ZONES 548
Push versus pull strategies 531 Endnotes 550
Pricing strategy 536

CHAPTER 13 INTERNATIONAL PRODUCTION,

OUTSOURCING AND LOGISTICS 552
OPENING CASE: LI & FUNG LI MITEO: Locating production facilities 567
A FACTORY OF SORTS 552 The Strategic Role of Foreign Factories 568
International Business Graduate Attributes (IBGA) 556 Outsourcing Production : Make-or-Buy Decisions 571
Learning Objectives 556 The advantages of 'make' decisions 572
Introduction 556 The advantages of 'buy' decisions 575
Strategy, Production and Logistics 558 Trade-offs 577
Where to Produce 560 Strategic alliances with suppliers 577
Country factors 560 Managing a Global Supply Chain 578
Technological factors 561 The role of just-in-time inventory 500
Product factors 566 Key Terms 583
 I CONTENTS I xiii

Summary 583 CLOSING CASE: FMCG, SOFT DOLLARS

International Business Graduate Attributes (IBGA): ANO LEAN SUPPLY CHAINS 586
Learning and Assessment Tasks 584 Endnotes 587

CHAPTER 14 INTERNATIONAL HUMAN

RESOURCE MANAGEMENT 590
OPEN ING CASE: ASTRAZENECA INTERNATIONAL 590 Petfonmance apl)(aisal problems 616
International Business Graduate Attributes (IBGA) 591 Guidelines for petformance appraisal 617
Learning Objectives 592 Compensation 618
Introduction 592 National differences in compensation 618
The Strategic Role of HRM 594 Expatriate pay 619
Staffing Policy 596 i nternationai Labour rleiations 623
Types of staffing policy 596 The concerns of organised labour 624
Expatriate management 601 Strategy of organised labour 625
Training and Management Development 610 Apl)(oaches to labour relations 628
Training for expatriate managers 611 Key Terms 628
Management development and strategy 613 Summary 629
Female Participation in Senior International International Business Graduate Attributes:
Management 613 Learning and Assessment Tasks 630
Equal opportunity 615 CLOSING CASE: CULTURAL NEGLECT AT THE
Work-life balance 615 MINGXI BING GUAN (GUEST HOUSE) 631
Mentorship and networking 616 Endnotes 634
Performance Appraisal 616

Glossary 638 Acronyms 659

Index 650 Countries/Capitals 663
PREFACE
Global Business Today is intended primarily as an • the 'emerging markets century'
introductory textbook for internationaVglobal business • concepts of global inequality
courses at universities at both the undergraduate and • the work of Nobel Prize-winning economist Amartya
postgraduate level. Sen on economic development
The authors have attempted to write a book that: • the work of Hernando de Soto on the link between
• presents a balanced global view property rights and economic development
• goes beyond an uncritical presentation and shallow • Samuel Huntington's influential thesis on the 'clash of
explanation of the body of knowledge civilisations'

• is well structured and well written, with tight • the new growth theory of economic development
integration and flow between chapters championed by Paul Romer and Gene Grossman
• focuses on managerial implications • empirical work by Jeffery Sachs and others on

• uses up-l~date examples and case studies the relation-ship between international trade and
• examines international business in contexts relevant economic growth
to the students' experiences • Michael Porter's theory on the competitive advantage
• makes important theories accessible and interesting of nations
• incorporates ancillary resources that enliven the text • Robert Reich's work on national competitive advantage
and make it easier to teach. • the work of Nobel Prize-winner Douglas North and
others on national institutional structures and the
In order to be comprehensive, a textbook on international
protection of property rights
business mus~
• Francis Fukuyama's proclamation of the 'end of
• explain how and why the world's countries differ history', by which global capitalism and democracy
• present a thorough review of the economics and would triumph around the world, and the critique
politics of international trade and investment arising from the Global Financial Crisis of 2007--09
• explain the functions and form of the international • Walden Bello's call for deglobalisation, the failure of
monetary system globalisation to deliver universal benefits, and the rise
• examine the strategies and structures of international of the economies of authoritarian China and Russia
businesses, both large and small • the 'market imperfections' approach to foreign direct
• examine the need for ethical and legal behaviour in investment that has grown out of Ronald Coase's and
international business Oliver Williamson's work on transaction cost economics
• assess the specific characteristics of international • Christopher Bartlett's and Sumantra Ghoshal's
business's various functions. research on the transnational corporation
As the authors of this book, we have endeavoured to do • the writings of C.K. Prahalad and Gary Hamel on core
all of these things. We have thoroughly examined the competencies, global competition and global strategic
structures and strategies of international firms, as well as alliances
the functional implications of international business for a • Pankaj Ghemawat's concept of semiglobalisation, and
variety of firms. Many students will soon be working in fields his work on how cross-border expansion is affected
that require an understanding of the effects of international by differences in the various dimensions of distance
business on an organisation's strategy, structure and • Makoto Kanda's work on long-lived firms
function. This book pays c lose attention to these issues. • insights into international business strategy that can
Comprehensiveness and relevance also require coverage be derived from the resource-based view of the firm.
of the major current theories. The following insights have
In light of the fast-changing nature of the international
therefore been included: business environment, the leading-edge theory and up-
• the new trade theory and strategic trade policy to-date material included in this book make ii a valuable
• theories of internationalisation, including 'born globals' resource in this field.
ABOUT THE AUTHORS
ABOUT THE ORIGINAL AUTHOR
CHARLES W. L. HILL is the Hughes M. Blake Professor of International Business at the School of Business, University
of Washington. Professor Hill received his PhD in industrial organisation economics in 1983 from the University of
Manchester's Institute of Science and Technology (UMIST) in Great Britain. In addition to his position at the University
of Washington, he has served on the faculties of UMIST, Texas A&M University and Michigan State University.
Professor Hill teaches in the MBA and executive MBA programs at the University of Washington and has received
awards for teaching excellence in both programs. He has also taught on several customised executive programs.
Professor Hill has published more than 50 articles in peer-reviewed academic journals, as well as four college
textbooks-one on strategic management, one on principles of management, and the other two on international business
(one of which you are now holding). He serves on the editorial boards of several academic journals and previously served
as consulting editor at the Academy of Management Review.

ABOUTTHELOCALAUTHORS
THOMAS CRONK was formerly an academic staff member of the School of International Business, Queensland
University of Technology (QUTJ. While in this school he led the ongoing development of the international business
curriculum in the Faculty of Business at QUT for nearly a decade. After retiring from full-time academic work, he served
until recently as a teaching and learning consultant to the Faculty of Business at QUT.
With more than 30 years' experience in business education in the higher education sector, Thomas has designed,
taught and administered a range of undergraduate and postgraduate courses, including Marketing and International
Business, Globalisation of Business, Global Business Operations and, for international students, Business in Australia.
He has also studied and developed courses in international business in several other countries, including the United
Kingdom, the United States, Malaysia and Hong Kong.
Thomas has authored other textbooks, including Economics for the Real World, Business Organisation and
Management and Marketing and International Business.

RUMINTHA WICKRAMASEKERA is the coordinator and senior lecturer of one of the largest International Business
programs in Australia-Global Business at Queensland University of Technology (QUTJ. He has more than 15 years'
experience in teaching, developing and coordinating international business subjects at a number of Australian universities.
Rumintha has also been involved in several major projects focusing on identifying the dynamics of the
internationalisation process of small to medium-sized enterprises, including his PhD research, which examined the
internationalisation of the Australian wine industry.
He is the recipient of several teaching awards including The QUT Vice-Chancellor's Award for Excellence in Learning
and Teaching, and the Australian Government Office for Learning and Teaching Citation for Outstanding Contributions to
Student Learning.
Another random document with
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spruce-pine forest by arranging these five formations in their proper
order in a succession herbarium, as follows: (1) the gravel slide
formation, (2) the half gravel slide formation, (3) the pine formation,
(4) the balsam-spruce formation, (5) the spruce-pine formation.

Development and Structure

247. Vegetation an organism. The plant formation is an

organic unit. It exhibits activities or changes which result in
development, structure, and reproduction. These changes are
progressive, or periodic, and, in some degree, rhythmic, and there
can be no objection to regarding them as functions of vegetation.
According to this point of view, the formation is a complex organism,
which possesses functions and structure, and passes through a cycle
of development similar to that of the plant. This concept may seem
strange at first, owing to the fact that the common understanding of
function and structure is based upon the individual plant alone.
Since the formation, like the plant, is subject to changes caused by
the habitat, and since these changes are recorded in its structure, it is
evident that the terms, function and structure, are as applicable to
the one as to the other. It is merely necessary to bear in mind that the
functions of plants and of formations are absolutely different
activities, which have no more in common than do the two
structures, leaf and zone.
248. Vegetation essentially dynamic. As an organism, the
formation is undergoing constant change. Constructive or
destructive forces are necessarily at work; the former, as in the plant,
predominate until maturity, when the latter prevail. Consequently, it
no longer seems fruitful to classify the phenomena of vegetation as
dynamic or static. The emphasis which has been placed upon
dynamic aspects of vegetation has served a useful purpose by calling
attention to the development of the latter. Although it is a quarter of
a century since Hult, and more than a half century since Steenstrup,
by far the greater number of ecological studies still ignore the
problem of development. This condition, however, can be remedied
more easily by insisting upon an exact understanding of the nature of
the formation than in any other way. It is entirely superfluous to
speak of dynamic and static effects in the plant, and the use of these
terms with reference to the formation becomes equally unnecessary
as soon as the latter is looked upon as an organism. The proper
investigation of a formation can no more overlook development than
structure, so closely are the two interwoven. Future research must
rest squarely upon this fact.
249. Functions and structures. The functions of a formation
are association, invasion, and succession: the second may be
resolved into migration and ecesis, and the third, perhaps, into
reaction and competition. Formational structures comprise zones,
layers, consocies, societies, etc., all of which may be referred to
zonation, or to alternation. The term association has been used in
both an active and a passive sense. In the former, it applies to the
inevitable grouping together of plants, by means of reproduction and
immobility. Passively, it refers to the actual groupings which result in
this way, and in this sense it is practically synonymous with
vegetation. Invasion is the function of movement, and of occupying
or taking possession; with association, it constitutes the two
fundamental activities of vegetation. It is the essential part of
succession, but the latter is so distinctive, because of the intimate
relation of competition and reaction, that clearness is gained by
treating it as a separate function which is especially concerned with
development. Association, zonation, and alternation are structural
phenomena, which are in large part the immediate product of habitat
and function, and in a considerable degree, also, the result of
ancestral or historical facts. It is a difficult matter to determine in
what measure the last factor enters, but it is one that must always be
taken into account, particularly when the physical factors of the
habitat are inadequate to explain the structures observed.
Structurally, association regularly includes both zonation and
alternation. As there are certain typical instances in which it exhibits
neither, the treatment will be clearer if each is considered separately.
 ASSOCIATION
250. Concept. The principle of association is the fundamental
law of vegetation. Indeed, association is vegetation, for the individual
passes into vegetation, strictly speaking, at the moment when other
individuals of the same kind or of different kinds become grouped
with it. It is then (and the same statement necessarily holds for
vegetation) the coming together and the staying together of
individuals and, ultimately, of species. A concrete instance will
illustrate this fact. In the development of the blowout formation of
the Nebraska sand-hills (Redfieldia-Muhlenbergia-anemium),
association begins only when the first plant of Redfieldia flexuosa is
joined by other plants that have sprung from it, or have wandered in
over the margin of the blowout. Henceforth, whatever changes the
blowout formation may undergo, association is a settled
characteristic of it until some new and overwhelming physical
catastrophe shall destroy the associated individuals. It will readily be
seen that association does not depend upon particular individuals,
for these pass and others take their place, but that it does depend
essentially upon number of individuals.
Association involves the idea of the relation of plants to the soil, as
well as that of plants to each other. It is synonymous with vegetation
only when the two relations are represented, since there may be
association such as that of a parasite with its host, which does not
constitute vegetation. But it will be seen that the relation of the
parasite to the host is practically identical with the relation of the
plant to the soil or stratum, and the two concepts mentioned above
become merged in such a case. From this it follows that association
results in vegetation only when the two ideas are distinct. The
concept of association contains a fact that is everywhere significant
of vegetation, namely, the likeness or unlikeness of the individuals
which are associated. In the case of parasite and host, this unlikeness
is marked; in vegetation, all degrees of similarity obtain. As will be
evident when the causes which lead to association are considered,
alternate similarity and dissimilarity of the constituent individuals or
species is subordinate as a feature of vegetation only to the primary
fact of association.
Since association contains two distinct, though related, ideas, it is
of necessity ambiguous. It is very desirable that this be avoided, in
order that each concept may be clearly delimited. For this reason, the
act or process of grouping individuals is termed aggregation, while
the word association is restricted to the condition or state of being
grouped together. In a word, aggregation is functional, association is
structural; the one is the result of the other. This distinction makes
clear the difference between association in the active and passive
sense, and falls in with the need of keeping function and structure in
the foreground.
251. Causes. In considering the causes which produce
association, it is necessary to call in evidence the primary facts of the
process in concrete examples of this principle. These facts are so
bound up in the nature of vegetal organisms that they are the veriest
axioms. Reproduction gives rise immediately to potential, and
ultimately, in the great majority of cases, to actual association. The
degree and permanence of the association are then determined by
the immobility of the individuals as expressed in terms of attachment
to each other or to the stratum, such as sheath, thallus, haustoria,
holdfasts, rhizoids, roots, etc. The range of immobility is very great.
In terrestrial plants, mobility is confined almost entirely to the
period when the individual lies dormant in the seed, spore, or
propagative part, which is alone mobile. In aquatic spermatophytes,
the same is true of all attached forms, while free floating plants such
as Lemna are mobile in a high degree, especially during the
vegetative period. Among the algae and hydrophilous fungi, attached
forms are mobile only in the spore or propagative condition, while
the motile forms of the plancton typify the extreme development of
mobility. The immediate result of reproduction in an immobile
species is to produce association of like individuals, while in the case
of a mobile species reproduction may or may not lead immediately to
association. We may lay down the general principle that immobility
tends to maintain the association of the individuals of the same
generation, i. e., the association of like forms, while mobility tends to
separate the similar individuals of one generation and to bring unlike
forms together. With the mobile algae, separation of the members of
each generation is the rule, unless the individuals come to be
associated in a thallus, or are grouped in contact with the
substratum. Flowering plants that are relatively immobile, especially
in the seed state, drop their seeds beneath and about the parent
plants, and in consequence dense association of the new plants is the
rule. In very many cases, however, this primitive tendency is largely
or completely negatived by the presence of special dissemination
contrivances, which are nearly, if not quite, as effective for many
terrestrial plants as the free floating habit is for algae. From this
point, the whole question of mobility belongs to migration, just as
the adjustment between the parent plants and their offspring, or
between plants established and the mobile plants to be established,
belongs to competition.
If association were determined by reproduction and immobility
alone, it would exhibit areas dissimilar in the mass of individuals, as
well as areas dissimilar in the kinds of individuals. Some areas would
be occupied by plants of a single species, others by plants of several
or many species. This tendency of association to show differences is,
however, greatly emphasized by the fact that vegetation is
fundamentally attached to and dependent upon a surface that
exhibits the most extreme physical differences. For this reason, new
differences in association appear, due not only to the morphological
differentiation of vegetation forms, but also to the changes in the
degree and manner of association produced directly by the different
habitats. Association might then be defined as a grouping together of
plant individuals, of parents and progeny, which is initiated by
reproduction and immobility, and determined by environment. It is a
resultant of differences and similarities. In consequence, association
in its largest expression, vegetation, is essentially heterogeneous,
while in those areas which possess physical or biological
definiteness, habitats and vegetation centers, it is relatively
homogeneous. This fundamental peculiarity has given us the concept
of the formation, an area of vegetation, or a particular association,
which is homogeneous within itself, and at the same time essentially
different from contiguous areas, though falling into a phylogenetic
series with some and a biological series with others. From its nature,
the plant formation is to be considered the logical unit of vegetation,
though it is not, of course, the simplest example of association.
 252. Aggregation. As indicated under the causes of association,
the process by which groups of individuals are formed depends
entirely upon reproduction and migration. In short, aggregation is
merely a corollary of movement. The simplest example of this
process occurs in forms like Gloeocapsa, Tetraspora, and others,
where the plants resulting from fission are held together by means of
a sheath. Though called a colony, such a group of individuals is a
family in the ordinary sense. Practically the same grouping results in
the case of terrestrial plants, especially spermatophytes, when the
seeds of a plant mature and fall to the ground about it. The relation
in both instances is essentially that of parent and offspring, although
the parent soon disappears in the case of annuals, while among the
algae its existence is regularly terminated by fission. The size and the
density of the family group are determined by the number of seeds
produced, and by their mobility. These are further affected by the
height and branching of the plant, and by the position of the seeds
upon it. The disseminules of immobile species fall directly beneath
the parent, and the resulting group is both uniform and definite. A
similar arrangement is caused likewise by offshoots. An increase in
mobility brings about a decrease of aggregation, since the
disseminules are carried away from the parent plant. Perfectly
mobile forms rarely produce family groups for this reason. It is
evident, however, that mobile perennials sometimes arrange
themselves in similar fashion in consequence of propagation by
underground parts. Consequently, it is possible to state the law of
single aggregation, viz., that immobility promotes the grouping of
parent and offspring, and mobility hinders it.
If all species were immobile, the family group would be
characteristic of vegetation. Since the great majority are more or less
mobile, aggregates of this sort are the exception rather than the rule.
Mobility not only decreases the number of offspring in the family
group, but it also spreads disseminules broadcast to enter dissimilar
groups. It leads directly to mixed aggregation, by which individuals
of one or more species invade the family group. Once established, the
newcomers tend also to produce simple groups, thus causing an
arrangement corresponding essentially to a community. Such
collections of family groups are extremely variable in size and
definition. This arises in part from the nature of simple aggregation,
and in part from the varying mobility of different species. Mobility
alone often produces similar communities by bringing together the
disseminules of different plants, each of which then becomes the
center of a mixed group. In the case of permobile species, several
disseminules of each may be brought together. The resulting area,
though larger, is practically the same. At present, it is difficult to
formulate the law for this method of grouping. It may be stated
provisionally as follows: mixed aggregation is the direct result of
mobility, and the greater the mobility the more heterogeneous the
mixture.
The constitution of all the major areas of a formation is to be
explained upon the basis of aggregation by the two methods
described. The relative importance of family groups and
communities differs for every formation, and the exact procedure in
each can be obtained only by the detailed study of quadrats. The
problem is further complicated by competition and reaction,
particularly in closed vegetation. For this reason, aggregation can be
studied most satisfactorily in a new or denuded area, where these
processes are not yet in evidence.

Kinds of Association

253. Categories. In the analysis of association, it must be kept

clearly in mind that the concrete examples from which all
generalizations must be drawn are often in very different stages of
development, and are of correspondingly different ages. For this
reason it has seemed best to consider the primary relations of
association in general in this place, leaving the treatment of the
effects of invasion, succession, alternation, and zonation to be taken
up under these topics.
Various categories of association may be distinguished, according
to the dominant physical factor concerned or the point of view taken.
These will fall into two series, as we consider the relation of plant to
plant with reference to some object or characteristic, or the grouping
of plants together in response to some dominant factor. In the first
series may be placed association with reference to substratum, to the
ground (occupation), and to invasion; in the second belong light and
water-content association. It should be noted that these are all actual
associations in nature, and not concepts such as the vegetation form,
within which plants from widely different associations may be
classified. Naturally, it does not follow that it is not logical or
valuable to group together those plants, such as hydrophytes,
sciophytes, hysterophytes, etc., which have a common relation to
some factor, but belong to different formations.
254. Stratum association. Plants manifest independent or
dependent association with reference to the stratum to which they
are attached and from which they derive food or support.
Independent association is exhibited by those holophytic species of a
formation which are entirely independent of each other with respect
to mechanical support or nutrition. It is characteristic of the greater
number of the constituent species of formations. Dependent
association is manifested in the relation between host and parasite,
stratum and epiphyte, support and liane. Warming[24] has
distinguished six kinds of associations: parasitism, helotism,
mutualism, epiphytism, lianism, and commensalism. Commensalism
corresponds to the primary principle of association which has given
rise to vegetation. Homogeneous commensalism is the term applied
to social exclusive plants, in which the patch is composed of a single
species. Such association is extremely rare in nature, and if the most
minute forms be considered, probably never occurs. On the other
hand, heterogeneous commensalism, in which individuals of more
than one species are present, is everywhere typical of vegetation.
Warming regards saprophytism merely as a specialized kind of
parasitism, an opinion that may well be defended. Helotism,
however, is also a mere modification of parasitism, if it is not indeed
parasitism pure and simple. Mutualism is an altogether vague
concept, including parasites, epiphytes, and endophytes of doubtful
physiological relation. Pound and Clements[25] treated lianes,
parasites, and saprophytes as vegetation forms, relating herbaceous
creepers and twiners to the lianes, and dividing the fungi and lichens
into nine groups. Whatever the value of these divisions may be from
the standpoint of vegetation forms, they represent the same relation
between plant and nutritive stratum, and with respect to association
should be merged in one group. Schimper[26] was the first to perceive
the essential similarity of all such groups from the standpoint of
association. He terms these plant societies (Genossenschaften),
retaining the four groups already established, lianae, epiphyta,
saprophyta, and parasiticae. It is evident that dependent association
comprises extremely divergent forms, from the slightly clinging herb,
such as Galium, to the most intense parasite. The distinction,
however, is a clear one, if restricted to that relation between plants in
which one acts as a mechanical support or stratum or as a nutritive
host for the other.
255. Ground association. The first division of formations into
open and closed was made by Engler and Drude.[27] Open formations
were defined as those having incomplete stability and heterogeneous
composition, while closed formations have a more definite uniform
stamp. What is true of formations is equally true of vegetation, so
that association may be regarded as open or closed with reference to
the density and thoroughness with which the plants occupy the
ground. In open association, the ground is slightly or partially
occupied, readily permitting the entrance of new plants without the
displacement of those already present. Such an arrangement is
characteristic of the early stages of a formation, or of a succession of
formations. It produces unstable open formations, which arise,
usually after denudation, in sand-hills, blowouts, gravel slides,
dunes, flood plains, burned areas, etc. In closed association,
occupation of the ground is complete, and the invasion of new
species can occur only through displacement. Closed association
results in stable, closed formations, such as forest, thicket, meadow,
and prairie. As open association characterizes the early stages of a
succession of formations, so closed association is peculiar to the later
or last stages of all such successions. In short, open formations
represent certain phases of the development of vegetation, while
closed formations correspond to the relatively final structural
conditions. It is a fundamental principle of association that every
succession from denudation, or from newly formed soils, begins with
open formations and ends with a closed formation. The causes
leading up to open and closed association are intimately connected
with development, and hence are considered under invasion and
succession.
256. Species guild association. Drude has distinguished a
kind of association peculiar to invasion, in which there is a successive
or concomitant movement of certain species of a formation into
another formation or region, resulting in species guilds
(Artengenossenschaften). The association in this case is largely one
of community of origin or area, and of concomitant migration. It is
especially characteristic of areas adjacent to formational and regional
limits. Fundamentally, it is merely the grouping of plants which are
invading at the same time, and consequently it differs only in degree
from what occurs in every invasion where more than a single
individual is concerned. Accordingly, this type of association has
little more than historical interest. This must not be construed to
mean that it does not occur, but that it differs in no essential from
the ordinary grouping of invaders.
257. Light association. The constituent species of formations
show two fundamentally different groupings with respect to light. In
the one case, the individuals are on the same level, or nearly so, in
such a way that each has direct access to sunlight. Such an
arrangement is characteristic of most grassland and herbaceous
formations. In the case of desert formations, there is often
considerable difference in the height of the plants, but the distance
between them is so great as to admit of direct illumination of all. This
arrangement may be termed coordinate association. In forests,
thickets, and many herbaceous wastes, the height and density of
certain species enable them to dominate the formation. In a dense
forest, the trees receive practically all the light incident upon the
formation, and the shrubs, herbs, fungi, and algae of lower habit and
inferior position must adapt themselves to the diffuse light which
passes through or between the leaves. The same is equally true of
dense thickets and wastes, except that the vertical distance is less,
and the diffuseness of the light is correspondingly modified. In these
formations, the dominant trees, shrubs, or herbs, the facies,
constitute a primary or superior layer. The degree of subordinate
association, as a result of which inferior layers will arise, is entirely
determined by the density of the facies. In open woodlands, which
are really mixed formations of woodland and grassland, the intervals,
and usually the spaces beneath the trees also, are covered with
poophytes, showing an absence of subordination due to light. This is
the prevailing condition in the pine formation (Pinus ponderosa-
xerohylium) of the ridges and foot-hills of western Nebraska. When,
however, the trees stand sufficiently close that their shadows meet or
overlap throughout the day, the increasing diffuseness begins to
cause modification and rearrangement of the individuals. By
photometric methods, the light in a forest is found to be least diffuse
just below the facies, while the diffuseness increases markedly in
passing to the ground. The taller, stronger individuals are
consequently in a position to assimilate more vigorously, and to
become still taller and stronger as a result. Just as these have taken
up a position inferior to that of the facies, so the shorter or weaker
species must come to occupy a still more subordinate position. This
results, not only because the light is primarily weaker nearer the
ground, but also because the taller plants interpose as a second
screen. The complete working out of this arrangement with reference
to light produces typical subordinate association, which finds its
characteristic expression in the layering of forests and thickets.
Layers tend to appear as soon as open woodland or thicket begins to
pass into denser conditions, and up to a certain point, at which they
disappear, they become the more numerous and the more marked,
the denser the forest.
In the Otowanie woods near Lincoln (Quercus-Hicoria-hylium),
layering usually begins at a light value of .1 (1 = normal sunshine in
the open). Thornber[28] has found the same value to obtain in the
thickets of the Missouri bluffs. In these, again, layers disappear at a
value of .005, the extreme diffuseness making assimilation
impossible except for occasional mosses and algae. A number of
herbaceous plants are present in the spring, but these are all
prevernal or vernal bloomers, which are safely past flowering before
shade conditions become extreme. In the Fraxinus-Catalpa-alsium,
all inferior holophytic vegetation disappears between the light value
of .004 and that of .003. The spruce-pine formation (Picea-Pinus-
hylium) of the Rocky mountains, with a light value of .01, usually
contains but a few scattered herbs, mostly evergreen; in some cases
there are no subordinate plants other than mosses and
hysterophytes. The lodge-pole pine formation (Pinus murrayana-
hylium), with light values often less than .005, is nearly or quite
destitute of all but hysterophytic undergrowth. Such extremely dense
formations are examples of coordinate association merely, since the
formation is reduced to a single superior layer, in which the
individuals of the facies bear the same spatial relation to incident
light. In layered formations, in addition to the subordinate relation
of other species to the facies, there is, of course, a kind of coordinate
association manifested in each layer.
 258. Water-content association. Schouw[29] was the first to
give definite expression to the value of the water-content of the soil
for the grouping of plants. He established four groups: (1) water
plants, (2) swamp plants, (3) plants of moist meadows, (4) plants of
dry soils. The first he termed hydrophytes, introducing the term
halophytes to include all saline plants. Thurmann[30] recognized the
fundamental influence of water-content upon association, and
further perceived that the amount of water present was determined
primarily by the physical nature of the soil. He distinguished plants
which grow in soils that retain water as hygrophilous, and those
found upon soils that lose water readily as xerophilous. Those which
seemed to grow indifferently upon either were termed ubiquitous.
The latter correspond in some measure to mesophytes, but they are
really plants possessing a considerable range of adaptability, and do
not properly constitute a natural group. Warming[31] proposed the
term mesophytes to include all the plants intermediate between
hydrophytes and xerophytes. He recognized the paramount value of
water-content association as the basis of ecology, and upon this
made a logical and systematic treatise out of the scattered results of
many workers. Schimper[32] placed the study of vegetation upon a
new basis by drawing a distinction between physical and
physiological water-content, and by pointing out that the last alone is
to be taken into account in the study of plant life, and hence of plant
geography. Accepting the easily demonstrable fact that an excess of
salts in the soil water, as well as cold, tends greatly to diminish the
available water of the soil, i. e., the chresard, it is at once seen why
saline and arctic plants are as truly xerophytic as those that grow on
rocks or in desert sands. An anomalous case which, however,
physical factor records have explained fully, is presented by many
plants growing in alpine gravel slides, strands, blowouts, sandbars,
etc., in which the water-content is considerable, but the water loss
excessive, on account of extreme heat or reduced air pressure. The
effect of these conditions is to produce a plant xerophytic as to its
aerial parts, and mesophytic or even hydrophytic as to subterranean
parts. Such plants may, from their twofold nature, be termed
dissophytes; they are especially characteristic of dysgeogenous soils
in alpine regions where transpiration reaches a maximum, but are
doubtless to be found in all gravel and sand habitats with high water-
content. With these corrections, the concept of water-content
association, which owes much to both Warming and Schimper, but is
largely to be credited to Thurmann, becomes completely and
fundamentally applicable to all vegetation.
Up to the present time, the general character of the habitat,
together with the gross appearance of the plant itself, has been
thought sufficient to determine the proper position of a plant or a
formation in the water-content classification. Such a method is
adequate, however, only for plants and formations which bear a
distinct impress. For an accurate classification into the three
categories, hydrophytes, mesophytes, and xerophytes, it is necessary
to make exact determinations of the normal holard and chresard of
the habitat, and to supplement this, in some degree at least, by
histological studies. Except in the case of saline, acid, and frozen
soils, the holard alone will be a fairly accurate index, especially in
habitats of similar soil composition. For an exact and comprehensive
classification, however, and particularly in comparative work, the
chresard must constitute the sole criterion. As the latter has been
ascertained for very few formations, and in Nebraska and Colorado
alone, the present characterization of many plants and formations as
hydrophytic, mesophytic, or xerophytic must be regarded as largely
tentative, and the final classification will be possible only after the
thorough quantitative investigation of their habitats.
The water-content groups, hydrophytia, mesophytia, and
xerophytia, include all formations found upon the globe. The
exactness with which this classification applies to vegetation is made
somewhat more evident by dividing mesophytia into forest and
grassland. This is based primarily upon light association, but it also
reflects water-content differences in a large degree. The groups thus
constituted represent the fundamental zonation of the vegetative
covering with respect to water-content. Ocean, forest, grassland, and
desert correspond exactly to hydrophytia, hylophytia, poophytia, and
xerophytia. The difference is merely one of terminology: the first
series takes into account the physiognomy of the vegetation itself,
while the other emphasizes the causative factors.
 THE DEVELOPMENT OF THE FORMATION
259. A strict account of development should trace the results of
the various activities of vegetation in their proper sequence. This is
aggregation, migration, ecesis, reaction, and competition. These
functions are so intimately and often so inextricably associated that
it is hardly feasible to discuss development by treating each one
separately. In consequence, the two fundamental phenomena,
invasion and succession, which they produce, are taken as the basis
of the discussion. These, moreover, are different only in degree;
succession is merely complete, periodic invasion. Nevertheless, the
subject gains in clearness by a separate treatment of each.
 INVASION
260. By invasion is understood the movement of plants from an
area of a certain character into one of a different character, and their
colonization in the latter. This movement may concern an individual,
a species, or a group of species. From the nature of invasion, which
contains the double idea of going into and taking possession of, it
usually operates between contiguous formations, but it also takes
place between formational zones and patches. More rarely and less
noticeably, there may be invasion into a remote vegetation, as a
result of long carriage by wind, water, birds, railroads, or vessels.
Movement or migration, however, represents but one of the two
ideas involved in invasion. Migration merely carries the spore, seed,
or propagule into the area to be invaded. In ecesis, the spores or
seeds germinate and grow, after more or less adjustment, and in case
the latter becomes sufficiently complete, the new plants reproduce
and finally become established. With all terrestrial plants, invasion is
possible only when migration is followed by ecesis, because of the
inherent differences of formations or of areas of the same formation.
In the case of surface floating forms, such as Lemnaceae, and of the
plancton, ecesis is of much less importance, on account of the
uniformity of the medium and the lack of attachment, and migration
is often practically synonymous with invasion.

MIGRATION

261. Migration has been sometimes used loosely as a synonym for

invasion, but it is here employed in its proper sense of removal or
departure, i. e., movement, and is contrasted with ecesis, the making
of a home, the two ideas being combined in invasion, which is a
moving into and a taking possession of. An analysis of migration
reveals the presence of four factors, mobility, agency, proximity, and
topography. Not all of these are present in every instance of
migration, as for example in the simple elongation of a rootstalk, but
in the great majority of cases each plays its proper part. Mobility
represents the inherent capacity of a plant for migration, and in its
highest expression, motility, is in itself productive of movement. As a
general rule, however, modifications for securing mobility are
ineffective in the absence of proper agents, and the effective
operation of the two will be profoundly influenced by distance and
topography.
262. Mobility denotes potentiality of migration as represented by
modifications for this purpose. It corresponds, in a sense, to
dissemination, though seed production also enters into it. Its most
perfect expression is found in those plants which are themselves
motile, Bacteriaceae, Oscillatoria, Volvocaceae, and Bacillariaceae,
or possess motile propagules, such as most Phycophyta. On the
other hand, it is entirely undeveloped in many plants with heavy
unspecialized seeds and fruits. Between these two extremes lie by far
the greater number of plants, exhibiting the most various degrees of
mobility, from the motile though almost immobile offshoots of many
Liliaceae to the immotile but very mobile spores of fungi. It is thus
seen that motility plays a relatively small part in migration, being
practically absent in terrestrial forms, and that it bears a very
uncertain relation to mobility. In analyzing the latter, contrivances
for dissemination are seen to determine primarily the degree of
mobility, while the number of seeds produced will have an important
effect in increasing or decreasing it. A third factor of considerable
importance is also involved, namely, position with reference to the
distributive agent, but any exact knowledge of its importance must
await systematic experiment somewhat after the methods of Dingler,
but with air-currents, etc., of known velocity and direction. The time
is not distant when by such methods it will be possible to establish a
coefficient of mobility, derived from terms of position, weight,
resistant surface, and trajectory for definite wind velocities or for
particular propulsive mechanisms.
263. Organs for dissemination. Plants exhibit considerable
diversity with reference to the part or organ modified, or at least
utilized, for dissemination. This modification, though usually
affecting the particular product of reproduction, may, in fact, operate
on any part of the plant, and in certain cases upon the entire plant
itself. In the majority of plants characterized by alternation of
generations, the same individual may be disseminated in one
generation by a reproductive body, and in the other by a propagative
one, as is the case in the oogones and conidia of Peronospora, the
spores and gemmae of Marchantia, the fruits and runners of
Fragaria, etc. Special modifications have, as a rule, been developed
in direct connection with spores and seeds, and mobility reaches its
highest expression in these. It is, on the other hand, greatly restricted
in offshoots and plant bodies, at least in terrestrial forms, though it
will now and then attain a marked development in these, as shown
by the rosettes of Sempervivum and the tumbling plants of
Cycloloma. For the sake of convenience, in analyzing migration, all
plants may be arranged in the following groups with reference to the
organ or part distributed.
1. Spore-distributed, sporostrotes. This includes all plants
possessing structures which go by the name of spore, such as the
acinetes of Nostoc and Protococcus, the zoogonidia of Ulothrix,
Ectocarpus, etc., the conidia, ascospores, and basidiospores of fungi,
the tetraspores of red seaweeds, and the gemmae and spores proper
of liverworts, mosses, and ferns. These are almost always without
especial contrivances for dissemination, but their extreme
minuteness results in great mobility.
2. Seed-distributed, spermatostrotes. This group comprises all
flowering plants in which the seed is the part modified or at least
disseminated. The mobility of seeds is relatively small, except in the
case of minute, winged or comate seeds.
3. Fruit-distributed, carpostrotes. The modifications of the fruit for
distribution exceed in number and variety all other modifications of
this sort. All achenes, perigynia, utricles, etc., properly belong here.
4. Offshoot-distributed, thallostrotes. To this class are referred
those plants, almost exclusively cormophytes, which produce lateral,
branch-like propagules, such as root-sprouts, rhizomes, runners,
stolons, rosettes, etc. Migration with such plants is extremely slow,
but correspondingly effective, since it is almost invariably followed
by ecesis.
5. Plant-distributed, phytostrotes. This group includes all plancton
and surface forms, whether motile or non-motile, and those
terrestrial plants in which the whole plant, or at least the aerial part,
is distributed, as in tumbleweeds and in many grasses.
 264. Contrivances for dissemination. Any investigation of
migration to be exact must confine itself to fixed forms. For these the
degree of perfection shown by dissemination contrivances
corresponds almost exactly to the degree of mobility. Because of the
difficulty of ascertaining the effect of ecesis, it is impossible to
determine the actual effectiveness in nature of different
modifications, and the best that can be done at present is to regard
mobility, together with the occurrence and forcefulness of
distributive agents, as an approximate measure of migration. The
general accuracy of such a measure will be more or less evident from
the following. Of 118 species common to the foot-hill and sand-hill
regions of Nebraska, regions which are sufficiently diverse to
indicate that these common species must have entered either one by
migration from the other, 83 exhibit modifications for
dissemination, while 8 others, though without special contrivances,
are readily distributed by water, and 4 more are mobile because of
minuteness of spore or seed. Some degree of mobility is present in 73
per cent of the species common to these regions, while of the total
number of species in which the mode of migration is evident, viz., 95,
66 per cent are wind-distributed, 20 per cent animal-distributed, and
14 per cent are water-distributed. It need hardly be noted that this
accords fully with the prevalence and forcefulness of winds in these
regions. Of the species peculiar to the foot-hill region, many are
doubtless indigenous, though a majority have come from the
montane regions to the westward. The number of mobile species is
121, or 60 per cent of the entire number, while the number of wind-
distributed ones is 85, or 70 per cent of those that are mobile. Among
the 25 species found in the widely separated wooded bluff and foot-
hill regions, 2 only, Amorpha nana and Roripa nasturtium, are
relatively immobile, but the minute seeds of the latter, however, are
readily distributed, and the former is altogether infrequent.
The following groups of plants may be distinguished according to
the character of the contrivance by which dissemination is secured:
1. Saccate, saccospores. Here are to be placed a variety of fruits, all
of which agree, however, in having a membranous envelope or an
impervious, air-containing pericarp. In Ostrya, Physalis, Staphylea,
the modification is for wind-distribution, while in Carex, Nymphaea,
etc., it is for water-transport.
 2. Winged, pterospores. This group includes all winged, margined,
and flattened fruits and seeds, such as are found in Acer, Betula,
Rumex, many Umbelliferae, Graminaceae, etc.
3. Comate, comospores. To this group belong those fruits and
seeds with long silky hairs, Gossypium, Anemone, Asclepias, etc.,
and those with straight capillary hairs or bristles not confined to one
end, Typha, Salix, etc.
4. Parachute, petasospores. The highly developed members of this
group, Taraxacum, Lactuca, and other Liguliflorae are connected
through Senecio and Eriophorum with the preceding. These
represent the highest development of mobility attained by special
modification.
5. Chaffy-pappose, carphospores. In this group are placed those
achenes with a more or less scaly or chaffy pappus with slight
mobility, as in Rudbeckia, Brauneria, Helianthus, etc.
6. Plumed, lophospores. In the fruits of this class, the style is the
part usually modified into a long plumose organ, possessing a high
degree of mobility, as in Pulsatilla, Sieversia, and Clematis.
7. Awned, ascospores. These are almost exclusively grasses, in
which the awns serve for distribution by wind, water, or animals, and
even, according to Kerner, by hygroscopic creeping movements. The
mobility in many cases is great.
8. Spiny, centrospores. This group contains a few representatives
which possess a moderate degree of mobility by attachment, as in
Tribulus and Cenchrus.
9. Hooked, oncospores. The members of this group are extremely
numerous, and the degree of mobility as a rule is very high. All
exhibit in common the development of hooks or barbs, by which they
are disseminated in consequence of attachment, though the number,
size, and disposition of the hooks vary exceedingly.
10. Viscid, gloeospores. In these, the inflorescence is more or less
covered with a viscid substance, as in species of Silene, or the fruit is
beset with glandular hairs, as in Cerastium, Salvia, etc.
11. Fleshy, sarcospores. These are intended for dissemination by
deglutition, largely by birds; the effectiveness of the modification
depends in a large degree upon the resistance of the seed envelope to
digestion. The mobility varies greatly, but the area over which
migration may be effected is large.
12. Nut-fruited, creatospores. This group includes those plants
with nut fruits which are carried away and secreted by animals for
food.
13. Flagellate, mastigospores. These are plants with ciliate or
flagellate propagative cells, i. e., zoogonidia, as in Protococcus,
Ulothrix, Oedogonium, Ectocarpus, etc., or with plant bodies
similarly motile, Bacteriaceae and Volvocaceae.
265. Position of disseminule. The position on the plant of the
organ to be disseminated, i. e., its exposure to the distributing agent,
plays a considerable part in determining the degree of mobility. In
the majority of plants, the position of the inflorescence itself results
in maximum exposure, but in a large number of forms special
modifications have been developed for placing the spores or seeds in
a more favorable position. In both cases, there are often present also
devices for bringing about the abscission of the seed or fruit. It is,
moreover, self-evident that the height of the inflorescence above
ground or above the surrounding vegetation is likewise of
considerable importance in increasing the trajectory. It is yet too
early to make a complete classification of contrivances for placing
disseminules in the most favorable exposure, but the following will
serve as a basis for future arrangements.
1. In all operculate Discomycetes, and especially in the
Ascobolaceae, where the asci project above the hymenium, the
spores are raised above the surface by tensions within the
apothecium. This might be regarded as dissemination by expulsion,
if it were not for the fact that the spores fall back into the cup, unless
carried away by the wind.
2. In Gasteromycetes and in certain Hepaticae, the spores are not
only elevated slightly above the sporophore by the expanding
capillitium or by the mass of elaters, but they are also held apart in
such a way that the wind blows them out much more readily.
3. In Bryophyta, the sporophore regularly dehisces by a slit, or is
provided with a peristome. Both structures are for the purpose of
sifting the spores out into the wind; by reason of their hygroscopicity,
they also insure that the spores will not be shaken out in wet
weather.
4. In a few grasses, such as Stipa and Aristida, the twisting and
intertwining of the awns lift the floret out of the glumes, and at the
same time constitute a contrivance readily blown away by the wind
or carried by attachment.
5. In certain Compositae, the involucral scales are reflexed at
maturity, and at the same time the disk becomes more or less
convex, serving to loosen the achenes. This result is also secured in
certain species by the drying and spreading of the pappus hairs.
6. The scapose Liguliflorae, Taraxacum, Agoseris, etc., are
characterized by the elongation of the scape after anthesis, with the
result that the head is raised to a considerable height by the time the
achenes are mature.
7. Carpotropic movements, though primarily for another purpose,
often serve to bring seeds and fruits into a better position for
dissemination.
266. Seed production. The relation of spore or seed-production
to mobility is obvious in the case of mobile species; in the case of
immobile ones, it is just as evident that it has no effect, though it may
still have considerable influence in increasing migration. In the case
of two species with equally effective dissemination contrivances, the
one with the largest seed-production will be the more mobile. On the
basis of the relation of seeds to flower, two groups of plants may be
distinguished, one, Polyanthae, in which the flowers are many and
the seeds few or single, as in Compositae, and the other,
Polyspermatae, Portulaca, Yucca, etc., in which the number of seeds
to each flower is large. So far as the actual number of seeds produced
is concerned, polyanthous plants may not differ from
polyspermatous ones, but, as a rule, they are much more highly
specialized for dissemination and are more mobile. The number of
fertile seeds is also much greater, a fact which is of great importance
in ecesis, and which, taken in connection with mobility, partially
explains the supremacy of the composites. Among the fungi and
algae, the amount of spore-production in a large degree determines
the mobility, since these forms are intrinsically permobile.
 267. Agents of migration. In the last analysis, however, the
possibility of migration depends upon the action of distributive
agents; in the absence of these, even the most perfect contrivance is
valueless, while their presence brings about the distribution of the
most immobile form. In short, migration depends much more upon
such agents than upon mobility, however perfect the latter may be. It
is, moreover, evident that the amount and extent of migration will be
determined primarily by the permanence and forcefulness of the
agent, as indicated by its ability to bring about transportation.
Finally, as will be shown later, the direction and rapidity of migration
depend directly upon the direction and intensity of the agent.
Migration results when spores, seeds, fruits, offshoots, or plants
are moved out of their home by water, wind, animals, man, gravity,
glaciers, growth, or mechanical propulsion. Corresponding to these
agents, there may be recognized the following groups:
1. Water, hydrochores. These comprise all plants distributed
exclusively by water, whether the latter acts as ocean currents, tides,
streams, or surface run-off. In the case of streams and run-off,
especially, mobility plays little part, provided the disseminules are
impervious or little subject to injury by water. Motile plants, or those
with motile cells, which belong entirely to this group, may be
distinguished as autochores, which correspond closely to
mastigospores.
2. Wind, anemochores. This group includes the majority of all
permobile terrestrial plants, i. e., those in which modifications for
increasing surface have been carried to the extreme, or those which
are already permobile by reason of the minuteness of the spore or
seed. Saccate, winged, comate, parachute, pappose, plumed, and, to a
certain extent, awned seeds and fruits represent the various types of
modifications for wind-distribution.
3. Animals, zoochores. Among terrestrial plants, dissemination by
attachment represents essentially the same degree of specialization
as is found in wind-distributed plants. The three types of
contrivances for this purpose are found in spinose, hooked, and
glandular fruits. Dissemination by deglutition and by carriage, either
intentional or unintentional, though of less value, play a striking part
on account of the great distance to which the seeds may be carried.
Dissemination by deglutition is characteristic of sarcospores, and
distribution by carriage of creatospores.
4. Man, brotochores. Dissemination by man has practically no
connection with mobility. It operates through great distances and
over immense areas as well as near at hand. It may be intentional, as
in the case of cultivated species, or unintentional, as in thousands of
native or exotic species. No other disseminating agent is comparable
with man in respect to universal and obvious migration.
5. Gravity, clitochores. The members of this group are exclusively
colline, montane, and alpine plants, growing on rocks, cliffs, and
gravel slides (talus), etc., in which the seeds reach lower positions
merely by falling, or more frequently by the breaking away and
rolling down of rock or soil masses and particles. Dissemination by
this method is relatively insignificant, though it plays an important
part in the rock fields and gravel slides of mountain regions,
particularly in the case of immobile species.
6. Glaciers, crystallochores. At the present time, transport by
glaciers is of slight importance, because of the restriction of the latter
to alpine and polar regions, where the flora is poorly developed. In
the consideration of migrations during the glacial epoch, however, it
plays an important point.
7. Growth, blastochores. The mobility of species disseminated by
offshoots is extremely slight, and the annual movement relatively
insignificant. The certainty of migration and of ecesis, is, however, so
great, and the presence of offshoots so generally the rule in
terrestrial plants that growth plays an important part in migration,
especially within formations.
8. Propulsion, bolochores. Like growth, dissemination by
mechanical propulsion, though operating through insignificant
distances, exerts an important effect in consequence of its
cumulative action. The number of plants, however, with contrivances
for propulsion is very much smaller than the number of blastochores.
All bolochorous species agree in having modifications by means of
which a tension is established. At maturity, this tension suddenly
overcomes the resistance of sporangium or fruit, and throws the
enclosed spores or seeds to some distance from the parent plant. In
accordance with the manner in which the tension is produced, sling-
fruits may be classified as follows:
 (a) Hygroscopicity, pladoboles. These include the ferns with
annulate sporangia, in which the expansion of the annulus by the
absorption of moisture bursts the sporangium more or less suddenly,
though the actual propulsion of the spores seems to come later as a
result of dessication.
(b) Turgescence, edoboles. Dissemination by turgescence is highly
developed in Pilobolus and in Discomycetes, though in the latter
turgescence results rather in placing the spores in a position to be
readily carried by the wind. Impatiens and Oxalis furnish familiar
examples of fruits which dehisce in consequence of increased
turgidity.
(c) Dessication, xerioboles. The number of fruits which dehisce
upon drying is very large, but only a small portion of these expel
their seeds forcibly. Geranium, Viola, Erysimum, and Lotus
illustrate the different ways in which dessication effects the sudden
splitting of fruits.
(d) Resilience, tonoboles. In some plants, especially composites,
labiates, and borages, the achenes or nutlets are so placed in the
persistent calyx or involucre that the latter serves as a sort of mortar
for projection, when the stem of the plant is bent to one side by any
force, such as the wind or an animal. It will be noticed that two
separate agents are actually concerned in dissemination of this sort.
Frequently, two or more agents will act upon the same
disseminule, usually in succession. The possibility of such
combinations in nature is large, but actual cases seem to be
infrequent, except where the activities of man enter into the
question. Some parts, moreover, such as awned inflorescences, are
carried almost equally well by wind or animals, and may often be
disseminated by the cooperation of these two agents. The wind also
often blows seeds and fruits into streams by which they are carried
away, but here again, parts adapted to wind-dissemination are
injured as a rule by immersion in water, and the number of plants
capable of being scattered by the successive action of wind and water
is small.
In the present state of our knowledge of migration, it is impossible
to establish any definite correspondence between dissemination-
contrivance, agent, and habitat. As a general rule, plants growing in
or near the water, in so far as they are modified for this purpose at
all, are adapted to water-carriage. Species which grow in exposed
grassy or barren habitats are for the most part anemochores, while
those that are found in the shelter of forests and thickets are usually
zoochorous, though the taller trees and shrubs, being exposed to the
upper air currents, are generally wind-distributed. There is then a
fair degree of correspondence, inasmuch as most hydrophytes are
hydrochorous, most hylophytes, zoochorous, and the majority of
poophytes and xerophytes, anemochorous. Definite conclusions can
be reached, however, only by the statistical study of representative
formations.
With respect to their activity, agents may be distinguished as
constant, as in the case of currents, streams, winds, slope, growth,
and propulsion, or intermittent, animals and man. In the former, the
direction is more or less determinate, and migration takes place year
by year, i. e., it is continuous, while in the latter dissemination is
largely an accidental affair, indeterminate in direction, and recurring
only at indefinite intervals. The effective conversion of migration into
invasion is greatest when the movement is continuous, and least
when it is discontinuous, since, in the latter, species are usually
carried not only out of their particular habitat but even far beyond
their geographical area, and the migration, instead of being an
annual one with the possibility of gradual adjustment, may not recur
for several years, or may, indeed, never take place again. The rapidity
of migration is greatest in the case of intermittent agents, while the
distance of migration is variable, being great chiefly in the case of
man, ocean currents, and wind, and slight when the movement is
due to slope, growth, or propulsion. Disregarding the great distances
over which artificial transport may operate, seeds may be carried half
way across the continent in a week by strong-flying birds, while the
possibilities of migration by growth or expulsion are limited to a few
inches, or at most to a few feet per year. This slowness, however, is
more than counterbalanced by the enormously greater number of
disseminules, and their much greater chance of becoming
established.
268. The direction of migration is determinate, except in the
case of those distributive agents which act constantly in the same
direction. The general tendency is, of course, forward, the lines of
movement radiating in all directions from the parent area. This is
well illustrated by the operation of winds which blow from any
quarter. In the case of the constant winds, migration takes a more or
less definite direction, the latter being determined to a large degree
by the fruiting period of any particular species. In this connection, it
must be kept clearly in mind that the position of new areas with
reference to the original home of a species does not necessarily
indicate the direction of migration, as the disseminules may have
been carried to numerous other places in which ecesis was
impossible. The local distribution of zoochorous species is of
necessity indeterminate, though distant migration follows the
pathways of migratory birds and animals. In so far as dissemination
by man takes place along great commercial routes, or along
highways, it is determinate. In ponds, lakes, and other bodies of
standing water, migration may occur in all directions, but in ocean
currents, streams, etc., the movement is determinate, except in the
case of motile species. The dissemination of plants by slopes,
glaciers, etc., is local and definite, while propulsion is in the highest
degree indeterminate. Migration by growth is equally indefinite, with
the exception that hydrotropism and chemotropism result in a
radiate movement away from the mass, while propulsion throws
seeds indifferently into or away from the species-mass. From the
above it will be seen that distant migration may take place by means
of water, wind, animals or man, and, since all these agents act in a
more or less definite direction over great distances, that it will be in
some degree determinate. On the other hand, local migration will as
regularly be indeterminate, except in the case of streams and slopes.
The direction of migration, then, is controlled by these distributive
agents, and the limit of migration is determined by the intensity and
duration of the agent, as well as by the character of the space through
which the latter operates.
 ECESIS

269. Concept. By the term ecesis is designated the series of

phenomena exhibited by an invading disseminule from the time it
enters a new formation until it becomes thoroughly established
there. In a word, ecesis is the adjustment of a plant to a new habitat.
It comprises the whole process covered more or less incompletely by
acclimatization, naturalization, accommodation, etc. It is the decisive
factor in invasion, inasmuch as migration is entirely ineffective
without it, and is of great value in indicating the presence and
direction of migration in a great number of species where the
disseminule is too minute to be detected or too little specialized to be
recognizable.
The relation of migration to ecesis is a most intimate one: the
latter depends in a large measure upon the time, direction, rapidity,
distance, and amount of migration. In addition, there is an essential
alternation between the two, inasmuch as migration is followed by
ecesis, and the latter then establishes a new center from which
further migration is possible, and so on. The time of year in which
fruits mature and distributive agents act has a marked influence
upon the establishment of a species. Disseminules designed to pass
through a resting period are often brought into conditions where
they germinate at once, and in which they perish because of
unfavorable physical factors, or because competing species are too
far advanced. On the other hand, spores and propagules designed for
immediate germination may be scattered abroad at a time when
conditions make growth impossible. The direction of movement is
decisive in that the seed or spore is carried into a habitat sufficiently
like that of the parent to secure establishment, or into one so
dissimilar that germination is impossible, or at least is not followed
by growth and reproduction. The rapidity and distance of migration
have little influence, except upon the less resistant disseminules,
conidia, gemmae, etc. Finally, the amount of migration, i. e., the
number of migrants, is of the very greatest importance, affecting
directly the chances that vigorous disseminules will be carried into
places where ecesis is possible.
 Normally, ecesis consists of three essential processes, germination,
growth, and reproduction. This is the rule among terrestrial plants,
in which migration regularly takes place by means of a resting part.
In free aquatic forms, however, the growing plant or part is usually
disseminated, and ecesis consists merely in being able to continue
growth and to insure reproduction. Here establishment is practically
certain, on account of the slight differences in aquatic habitats,
excepting of course the extremes, fresh water and salt water. The
ease indeed with which migration and ecesis are effected in the water
often makes it impossible to speak properly of invasion in this
connection, since aquatics are to such a large extent cosmopolitan. In
dissemination by offshoots, the conditions are somewhat similar.
Here, also, ecesis comprises the sequence of growth and
reproduction, and invasion, in the sense of passing from one habitat
to another, is of rare occurrence, as the offshoot grows regularly
under the same conditions as the parent plant. The adjustment of
growing plants and parts is so slight, and their establishment so
certain on account of their inability to migrate into very remote or
different habitats, that they may be ignored in the following
discussion.
In accordance with the above, it would be possible to distinguish
three groups of terrestrial plants: (1) those migrants which germinate
and disappear, (2) those which germinate and grow but never
reproduce, (3) those which reproduce, either by propagation or
generation, or both. Such a classification has little value, however,
since the same species may behave in all three fashions, depending
upon the habitat to which it has migrated, and since invasion does
not occur unless the plant actually takes possession, i. e., reproduces.
From the latter statement, it follows that invasion occurs only when a
species migrates to a new place, in which it germinates, matures, and
reproduces. Maintenance by annual invasion simply, in which the
plants of each year disappear completely, can not then be regarded
as invasion proper. On the other hand, though such instances are
rare, it is not necessary that the invaders produce fruit, provided they
are able to maintain themselves, or to increase by propagation.
Furthermore, if a plant germinate, grow, and reproduce, it is
relatively immaterial whether it persist for a few years or for many,
since, as we shall see under Succession, the plants of one invasion are
displaced by those of the next, the interval between invasions
increasing with the stabilization.
270. Germination of the seed. The germination of seed or
spore is determined by its viability and by the nature of the habitat.
Viability depends upon the structural characters of fruit, seed-coat,
and endosperm, and to a degree upon the nature of the protoplasm
or embryo. The first three affect the last directly, by protecting the
embryo against dryness, against injury due to carriage by water, or
by deglutition, and probably in some cases against excessive heat or
cold. Marloth[33] has investigated the structure of seed coats,
establishing the following groups, which are summarized here
somewhat fully because of their bearing upon ecesis: (1) seed coats
without protective elements, endosperm absent or rudimentary,
Epilobium, Impatiens, Parnassia, Sagittaria, etc.; (2) protective
elements lacking or few, endosperm highly developed with thick-
walled cells, Liliaceae, Primulaceae, Rubiaceae, etc.; (3) protective
cells present in the seed coats, endosperm little or none,
Boraginaceae, Crassulaceae, Cruciferae, Labiatae, Papilionaceae,
etc.; (4) protective elements present, Asclepias, Campanula,
Gentiana, Silene, Saxifraga, etc.; (5) protective cells present,
endosperm thick-walled, Euonymus, Helianthemum, Ribes. The
protective cells are of various kinds: (1) epidermal cells strongly
cuticularized, Caryophyllaceae, Crassulaceae, Fumariaceae,
Saxifragaceae; (2) parenchyma thick-walled, several-layered,
Aesculus, Castanea, Fagus; (3) parenchyma cells with the inner or
radial walls thickened, Campanula, Erythraea, Gentiana; (4)
epidermal cells cup-shaped, thick-walled, Cruciferae, Ribes,
Vaccinium; (5) parenchyma with thickened, cellulose walls,
Geranium, Viburnum; (6) a single row of stone-cells, Labiatae; (7)
tissue of stone-cells, Hippuris, Naias, Potamogeton; (8) elongate
stone-cells, Coniferae, Cupuliferae, Euphorbia, Linum, Malva,
Viola; (9) short, columnar, thick-walled branched cells,
Cucurbitaceae, Datura, Hypericum; (10) prosenchyma with
cellulose walls, Clematis; (11) prosenchyma with lignified walls,
Fraxinus, Rhamnus, Ranunculus. The seed coats have a certain
influence in determining germination at the proper time, inasmuch
as they make it difficult for the seed to germinate under the stimulus
of a quantity of warmth and moisture insufficient to support the
seedling. The effect of the endosperm, as well as that of other food
supply in the seed, upon germination and the establishment of the
seedling is obvious.
The behavior of seed or spore with respect to germination depends
in a large degree upon the character of the protoplasm or embryo,
though in just what way is at present a matter of conjecture. It is
evident that many seeds are not viable because fertilization has not
been effected, and in consequence no embryo has developed. This is
the usual explanation of the low germinating power of the seeds of
some species, especially polyspermatous ones. But even in viable
seeds the behavior is always more or less irregular. The seeds of
some species will grow immediately after ripening, while others
germinate only after a resting period of uncertain duration. The same
is true of spores. Even in the case of seeds from the same parent,
under apparently similar conditions, while the majority will
germinate the first year, some will lie dormant for one or more years.
The precise reason why many seeds and spores germinate more
readily after being frozen is equally obscure. The period of time for
which disseminules may remain viable is extremely diverse, though,
as would be expected, it is much longer as a rule for seeds than for
spores. The greater vitality of seeds in the case of ruderal plants
suggests that this diversity may be due simply to variation in the
vigor of the embryos. It would seem that under proper conditions
seeds may retain their viability for an indefinite period.
The influence of habitat upon germination is of primary
importance, though the manner in which its influence is exerted is by
no means as evident as might be supposed. In the case of seeds sown
in the planthouse, it is almost universally the case that germination
is less than in nature, notwithstanding the fact that temperature and
moisture appear to be optimum. In nature, the seeds of the species
may be carried into a number of different formations, any one or all
of which may present conditions unfavorable to germination. With
respect to probability of germination, habitats are of two sorts: those
which are denuded and those which bear vegetation. It is impossible
to lay down general propositions with respect to either group, since
germination will vary with the character of the invading species, the
annual distribution of heat and moisture in the habitat, etc. In a
general way, however, it may be stated that the chances for
germination are greater in vegetation than in denuded areas, chiefly