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Preface

The transformation in marketing research practiced in organizations the world over is telling.
Across a multitude of data sources and technology, marketers are absorbing the developments at
a rapid pace. This warrants a need to be constantly up-to-date with the knowledge and capabilities
to channel these developments to achieve marketing and financial success. The nature and type
of data being collected now also include not only store scanner data, but also consumer home
shopping panels and data from social media. The rapid expansion of social media outlets such as
blogs, forums, and user networks has increasingly taken a more primary role among marketers.
With respect to word-of-mouth, product information is routinely shared between and across users
and nonusers that has the potential to create new customers. Now more than ever companies have
the ability to collect a wide range of customer information pertaining to attitudes, behaviors, and
demographics. An expansion in data storage and retrieval techniques (ranging from small data-
bases to huge data farms), coupled with declining data storage costs, has propelled the growth
of marketing research by leaps and bounds over the past decades. These developments have
enabled companies to invest in data now more than ever. In recent years, technological develop-
ments such as Internet of Things, artificial intelligence, and blockchain continue to dictate the
course of business practices. The promise of customer-level interactions for firms adopting such
technological developments is astounding. Alongside this promise comes the issue of privacy. In
these formative years of new technological developments, consumers and firms are trying to find
a middle ground with respect to data exchange and data privacy. Balancing such new develop-
ments with traditional know-how in marketing research is the new challenge for marketers. This
new edition of Marketing Research brings to the forefront the relevance of marketing intelligence
amidst these changes.
If we can compare marketing to a long train with multiple compartments, then marketing
research would justly claim the dual roles of the engine that powers the train and the links that
connect the individual compartments to form a cohesive functional unit. In other words, market-
ing research is pervasive—the brain and the brawn of any marketing organization. Having said
this, we realize that marketing research is a complex subject and therefore has to be introduced
to the student one compartment at a time before the entire train can be visualized. We also realize
the danger in this approach. The student can get overly excited or, even worse, overwhelmed by
the individual units so that he or she fails to see the proverbial “big picture”—the overarching
framework, the subtle but essential interactions between units, and the ultimate purpose, namely,
how marketing research can help organizations achieve their goals.
This revised edition takes a “macro–micro–macro” approach toward communicating the
intricacies of marketing research and its usefulness to the marketing organization. The first two
chapters provide an overview of the marketing research process that illustrates the “big picture.”
The chapters that follow then discuss in detail the individual components. The chapters have
benefitted significantly from updated business examples and industry statistics. While main-
taining the strengths of previous editions, this edition focuses on the recent trends in marketing
intelligence. Specific importance has been given to the concepts such as Customer Lifetime
Value, Share of Wallet, Brand Equity, Mobile Marketing, and the increasing role of Social Media

vii
viii Preface

Marketing. Throughout the book, examples of companies using technology developments such
as bots and big data have been introduced to inform the student about how fundamental topics
in marketing research have been applied in the new technology-focused environment. Topics
of less interest and relevance to the practice of marketing and marketing research have been
eliminated.
We begin with a macro-level treatment of what marketing research is, where it fits within
an organization, and how it helps in managerial decision making. Here, we also discuss the
marketing research industry, with a brief treatment of both suppliers and users.
The body of the text takes a micro-level approach, detailing each and every step of the market-
ing research process. In describing the marketing research process, a decision-oriented perspec-
tive has been adopted to help students, who are future managers and researchers, make better
decisions. Detailed discussions of the process, with numerous examples from the industry, char-
acterize this micro phase.
Finally, we wrap up with a macro-level treatment of the applications of marketing research.
Here we address the traditional 4P research, as well as contemporary issues such as brand equity,
customer satisfaction research, and emerging issues that continue to fascinate marketers, such as
e-commerce, direct marketing, database marketing, and relationship marketing, while taking care
to incorporate some of the latest research and developments in these fields.

Objectives of this Text

Our overall objectives in writing this text continue to be the following:
1. To emphasize the role of marketing research in today’s world and to focus on the tech-
niques and steps that show how a company can gather marketing intelligence and use this
information to generate insights that can be used in strategic decision making.
2. To communicate in an interesting and informative manner the essence of marketing
research to “future managers” and “future researchers.” Both groups need to know when
marketing research can and should be used, what research alternatives exist, how to recog-
nize effective and ineffective research, and how to interpret and apply the results.
3. To illustrate the usefulness of the Internet, online marketing research, and other advances
in technology in collecting data and show real-world applications.
4. To emphasize the current developments in marketing research, such as the distinction
between domestic and international market research, the emphasis on customer value man-
agement, and the influence of social media.
5. To use examples, applications, and illustrations throughout the book, in an effort to tie
the material to the real world and thus provide interest and better understanding to the
student.
6. To discuss the fastest growing applications of marketing research, e-commerce, direct mar-
keting, database marketing research, customer relationship management, and social net-
working and their impact on businesses.
7. To provide a clear and comprehensive treatment of data analysis topics. Each chapter
includes simple numerical examples to help students get a hands-on feel for the material.
8. To provide a thorough coverage of the most advanced and current marketing research meth-
odologies, pointing out their limitations as well as their potential for enhancing research
results.
 Preface ix

New to This Edition

Consistent with these objectives, the 13th edition has undergone some critical changes. The more
prominent of these include the following:
1. The chapter objectives have been clearly detailed on the first page of each chapter in order
to clearly communicate what is to be covered in the chapter.
2. Chapter 1 (A Decision-Making Perspective on Marketing Intelligence) has been updated
to cover the latest business trends and company examples that aptly reflect the nature and
level of changes made throughout the textbook.
3. Chapter 2 (Marketing Research in Practice) offers an updated report of the marketing
research industry in the United States and around the world.
4. Chapter 6 (Standardized Sources of Marketing Data) contains updated information on
marketing data sources that accounts for the rapid changes in technology that impacts
data collection. Also, this chapter lists examples of companies using such modes of data
sources.
5. Chapter 7 (Marketing Research on the Internet) contains revised and updated informa-
tion regarding the Internet and how it impacts marketing research. Also, the promi-
nence of privacy and safety issues on the Internet have been elaborated using recent
developments.
6. Chapter 25 (New Age Strategies) has been significantly revised and updated. Reflecting
the chapter title, this chapter has undergone changes that reflect the latest trending market-
ing strategies. Sections that are outdated have been removed, and new sections have been
added. Also, new topics such as Internet of Things, big data, and blockchain have been
covered in this edition.
7. The examples contained in the Marketing Research in Action throughout the book have
been updated with relevant content changes.
8. Statistics and other trend information have been updated. Further, new cases and study
questions have been added.
9. The text has also benefitted from additions and updates to new and emerging marketing
topics such as lifetime value, and measuring the effectiveness of social media campaigns,
and technology topics such as artificial intelligence and bots.
10. The Web address www.wiley.com/go/Kumar/marketingresearch13e can be used for access-
ing information pertaining to the textbook and its contents.

Features of the Book

1. The text communicates the essence of marketing research in an interesting and informative
manner to future managers and future researchers. Both groups need to know when mar-
keting research can and should be used, what research alternatives exist, how to recognize
effective and ineffective research, and how to interpret and apply the results.
2. The Cases and Part Cases are positioned appropriately at the end of chapters and text parts
to stimulate interest, add realism to the marketing research curriculum, and help develop
decision-making skills. These cases cover a wide range of products and organizations.
x Preface

3. The chapter on A Decision-Making Perspective on Marketing Intelligence attempts to

enlighten the readers of the importance and means of accessing data from multiple sources
and delivering to end-users for analysis.
4. The chapter on Marketing Research on the Internet links the reader with the world of
marketing research to keep abreast with the emerging trends and changes in the market-
place.
5. Discussion of the international element of marketing research has been continued. Par-
ticularly, an effort has been made to provide a clear distinction between the domestic and
international marketing research processes and prepare the users of this text to face the
challenges of multinational research.
6. The chapter on New Age Strategies reiterates the fact that understanding of the customer
is the key to marketing success. The chapter focuses on the tools provided by marketing
research vis-à-vis the concepts and technology. Whereas the developments in marketing
concepts such as database marketing and relationship marketing are discussed, the techno-
logical developments such as Internet of Things, bots, artificial intelligence, and blockchain
are also covered. The balance in attention between marketing concepts and technology
developments covered in this chapter aptly reflects the dynamics currently playing in the
market. As firms shift their resources more toward targeted marketing, the discussion in
this chapter becomes more valuable.
7. Marketing Research in Action sections in various chapters have been updated to focus on
the real-world applications of Marketing Research.
8. Each chapter also includes Learning Objectives, a Chapter Summary, and Questions and
Problems.

Organization of the Text

The book is organized to reflect the “macro–micro–macro” approach toward imparting market-
ing research training to the student. The text consists of five parts. Parts I and V deal with the
“macro” aspects of marketing research; Parts II–IV deal with the micro aspects.
Part I, consisting of four chapters, deals with the nature and scope of marketing intelligence
and marketing research. Here, the overall framework of marketing research is presented, and
where and how marketing research fits in with the other aspects of marketing is explained.
The nature of the research industry and suppliers is also discussed here.
Part II, consisting of Chapters 5 through 15, deals extensively with the various aspects of data
collection. This part is further divided into four sections, one section devoted to each of the
three fundamental types of marketing research: exploratory, descriptive, and causal. The final
section addresses the issue of sampling.
Part III, consisting of three chapters, discusses the fundamental aspects and techniques in
data analysis. These include basic analysis issues such as data editing, coding, and simple
techniques such as hypothesis testing, chi-square analysis, and the analysis of variance.
Part IV is devoted exclusively to advanced and more sophisticated data analysis techniques
such as correlation and regression analysis, discriminant analysis, factor analysis, cluster
analysis, conjoint analysis, and multidimensional scaling. This part consists of four chapters,
the last chapter dealing with the aspects of presenting the research results.
 Preface xi

In Part V, the student is exposed to the measures, metrics, and strategies of marketing intelli-
gence. This section has three chapters and provides the student with a comprehensive picture
of marketing research, highlighting how the marketing-mix measures, the brand and cus-
tomer metrics, and the new age strategies fit into the application of marketing intelligence.

Supplements to the Text

• All relevant information pertaining to the textbook and its contents can be accessed from
this website—www.wiley.com/go/Kumar/marketingresearch13e. This website will periodi-
cally update information that is relevant for keeping the text up to date.
• An Instructor’s Manual will accompany this text. This manual provides solutions to
end-of-chapter Questions and Problems and discusses all the test cases in greater detail.
Exam questions are arranged by chapter and include multiple-choice and true/false ques-
tions. An example of a course syllabus is presented, and many suggestions for the organiza-
tion of the course are provided. This resource can be found on the Instructor Companion site.
• A test bank has been added to the website, which provides sample test questions in each
chapter.
• An Image Gallery with figures for each chapter is available.
• Data sets that contain data for end-of-chapter cases and statistical analysis are available.
SPSS®-interpreted examples are also available.
• The website also contains detailed information on the mini-project designed for students
mentioned at the end of the eighth chapter.
• End-of-chapter questions are also available on the website.

Acknowledgments
Many debts have been accumulated over the years during which 13 editions of this book have
taken shape. We are especially grateful to our students, who gave us feedback from the con-
sumer’s perspective and whose field research projects provided many of the illustrations and
problems; to our colleagues, who stimulated us and brought new ideas and approaches to our
attention; and to our clients, who gave us many opportunities to put ideas into practice and
thus broaden our understanding of marketing research as it is currently practiced. It has been a
continuing pleasure to associate with a class publisher, John Wiley & Sons, and to work with a
number of Wiley editors over the years—.
We also would like to express thanks to Bharath Rajan and Divya Ramachandran for their
efforts toward the production of the 13th edition.
Finally, we wish to express our sincere appreciation to our families and friends for their con-
stant support, encouragement, and sacrifices during the creation of this book.

V. Kumar
Robert P. Leone
David A. Aaker
George S. Day
Brief Contents

Part I The Nature and Scope of Marketing Research

1 A Decision-Making Perspective on Marketing Intelligence 1
2 Marketing Research in Practice 30
3 The Marketing Research Process 47
4 Research Design and Implementation 71

Part II Data Collection

Section A: Secondary and Exploratory Research

5 Secondary Sources of Marketing Data 98

6 Standardized Sources of Marketing Data 123
7 Marketing Research on the Internet 147
8 Information Collection: Qualitative and Observational Methods 173

Section B: Descriptive Research

9 Information from Respondents: Issues in Data Collection 209
10 Information from Respondents: Survey Methods 226
11 Attitude Measurement 255
12 Designing the Questionnaire 284

Section C: Causal Research

13 Experimentation 313

Section D: Sampling
14 Sampling Fundamentals 344
15 Sample Size and Statistical Theory 372

xiii
xiv Brief Contents

Part III Data Analysis

16 Fundamentals of Data Analysis 396
17 Hypothesis Testing: Basic Concepts and Tests of Associations 416
18 Hypothesis Testing: Means and Proportions 435

Part IV Special Topics in Data Analysis

19 Correlation Analysis and Regression Analysis 461
20 Discriminant, Factor, and Cluster Analysis 491
21 Multidimensional Scaling and Conjoint Analysis 533
22 Presenting the Results 556

Part V Applications of Marketing Intelligence

23 Marketing-Mix Measures 571
24 Brand and Customer Metrics 613
25 New Age Strategies 635
Appendix A 673
Glossary 699
Index 717
Contents

Part I The Nature and Scope of Marketing Research

1 A Decision-Making Perspective on Marketing Intelligence 1

Learning Objectives, 1
An Overview of Business Intelligence, 1
Introduction to Marketing Intelligence, 4
Marketing Research, 7
Role of Marketing Research in Managerial Decision-Making, 9
Factors that Influence Marketing Research Decisions, 17
Use of Marketing Research, 19
Ethics in Marketing Research, 20
The Respondent’s Ethics and Rights, 21
International Marketing Research, 23
Summary, 25
Questions and Problems, 25
End Notes, 26
Case 1-1: Preteen Market—The Right Place to Be In for Cell Phone
Providers?, 27
Case 1-2: Best Buy on a Segmentation Spree, 27
Case 1-3: Ethical Dilemmas in Marketing Research, 29

2 Marketing Research in Practice 30

Learning Objectives, 30
Information Systems, Decision Support Systems, and Marketing Research, 31
Marketing Decision Support Systems, 34
Suppliers of Information, 37
Criteria for Selecting External Suppliers, 43
Career Opportunities in Marketing Research, 44
Summary, 45
Questions and Problems, 45
End Notes, 45
Appendix: Marketing Research Jobs, 46

3 The Marketing Research Process 47

Learning Objectives, 47
Overview of the Marketing Research Process, 47

xv
xvi Contents

The Preliminary Stages of the Marketing Research Process, 48

Planning a New HMO, 60
The International Marketing Research Process, 62
Summary, 66
Questions and Problems, 66
End Notes, 67
Case 3-1: A VideOcart Test for Bestway Stores, 67
Case 3-2: Philip Morris Enters Turkey, 69

4 Research Design and Implementation 71

Learning Objectives, 71
Research Approach, 73
Research Tactics and Implementation, 79
Budgeting and Scheduling the Research Project, 81
Research Proposal, 83
Designing International Marketing Research, 84
Issues in International Research Design, 86
Summary, 88
Questions and Problems, 89
End Notes, 89
Appendix: Errors in Research Design, 90
Case 4-1: Reynolds Tobacco’s Slide-Box Cigarettes, 91
Cases for Part I, 95
Case I-1: Clover Valley Dairy Company, 95

Part II Data Collection

Section A: Secondary and Exploratory Research

5 Secondary Sources of Marketing Data 98

Learning Objectives, 98
Secondary Data, 99
Uses of Secondary Data, 100
Benefits of Secondary Data, 100
Limitations of Secondary Data, 101
Internal Sources of Secondary Data, 102
External Sources of Secondary Data, 103
Census Data, 109
North American Industry Classification System, 111
Appraising Secondary Sources, 112
Applications of Secondary Data, 113
Sources of Secondary Data for International Marketing Research, 117
Problems Associated with Secondary Data in International Research, 117
 Contents xvii

Applications of Secondary Data in International Research, 118

Summary, 119
Questions and Problems, 119
End Notes, 120
Case 5-1: Barkley Foods, 121
Case 5-2: Dell in Latin America?, 122

6 Standardized Sources of Marketing Data 123

Learning Objectives, 123
Retail Store Audits, 124
Consumer Purchase Panels, 125
Scanner Services and Single-Source Systems, 128
Media-Related Standardized Sources, 134
Applications of Standardized Sources of Data, 136
Summary, 138
Questions and Problems, 138
End Notes, 139
Case 6-1: Promotion of Rocket Soups, 140
Case 6-2: Kerry Gold Products, Ltd., 144

7 Marketing Research on the Internet 147

Learning Objectives, 147
Current Trends in Web Usage, 148
WWW Information for Marketing Decisions, 149
The Internet and Marketing Research Today, 154
The Internet and Marketing Research Developments, 166
Issues and Concerns, 167
Summary, 169
Questions and Problems, 169
End Notes, 170
Case 7-1: Caring Children’s hospital, 172

8 Information Collection: Qualitative and Observational

Methods 173
Learning Objectives, 173
Need for Qualitative Research, 173
Qualitative Research Methods, 174
Observational Methods, 194
Recent Applications of Qualitative and Observational Methods, 200
Summary, 201
Questions and Problems, 201
End Notes, 202
Appendix: Myths of Qualitative Research: It’s Conventional, but Is It
Wisdom?, 204
xviii Contents

Case 8-1: Mountain Bell Telephone Company, 205

Case 8-2: U.S. Department of Energy (A), 207
Case 8-3: Hamilton Beach Conducts Primary Research in
Mexico and Europe, 208

Section B: Descriptive Research

9 Information from Respondents: Issues in Data Collection 209

Learning Objectives, 209
Information from Surveys, 209
Sources of Survey Error, 210
Methods of Data Collection, 215
Factors Affecting the Choice of a Survey Method, 216
Ethical Issues in Data Collection, 220
Summary, 222
Questions and Problems, 222
End Notes, 223
Case 9-1: Essex Markets, 224
Case 9-2: More Ethical Dilemmas in Marketing Research, 224

10 Information from Respondents: Survey Methods 226

Learning Objectives, 226
Collecting Data, 226
Surveys in the International Context, 248
Summary, 250
Questions and Problems, 250
End Notes, 252
Case 10-1: Roland Development Corp, 253

11 Attitude Measurement 255

Learning Objectives, 255
Attitudes, 256
The Concept of Measurement and Scaling, 257
Types of Attitude Rating Scales, 260
General Guidelines for Developing a Multiple-Item Scale, 271
Interpreting Attitude Scales, 274
Choosing an Attitudinal Scale, 275
Accuracy of Attitude Measurements, 277
Scales in Cross-National Research, 279
Summary, 280
Questions and Problems, 280
End Notes, 281
Case 11-1: National Kitchens, 282
 Contents xix

12 Designing the Questionnaire 284

Learning Objectives, 284
Planning What to Measure, 285
Formatting the Question, 287
Question Wording: A Problem of Communication, 292
Sequence and Layout Decisions, 297
Questionnaire Design for International Research, 303
Summary, 305
Questions and Problems, 305
End Notes, 309
Case 12-1: Wine Horizons, 309
Case 12-2: Smith’s Clothing (A), 312

Section C: Causal Research

13 Experimentation 313
Learning Objectives, 313
Descriptive versus Experimental Research, 314
What Constitutes Causality?, 315
Laboratory and Field Experiments, 317
Threats to Experimental Validity, 318
Issues in Experimental Research, 320
Types of Experimental Designs, 322
Guidelines for Conducting Experimental Research, 336
Limitations of Experiments 337
Summary, 339
Questions and Problems, 339
End Notes, 341
Case 13-1: Evaluating Experimental Designs, 341
Case 13-2: Barrie Food Corporation, 342

Section D: Sampling

14 Sampling Fundamentals 344

Learning Objectives, 344
Sample or Census, 344
Sampling Process, 346
Probability Sampling, 352
Nonprobability Sampling, 357
Shopping Center Sampling, 362
Sampling in the International Context, 363
Summary, 365
Questions and Problems, 366
xx Contents

End Notes, 368

Case 14-1: Exercises in Sample Design, 369
Case 14-2: Talbot Razor Products Company, 369

15 Sample Size and Statistical Theory 372

Learning Objectives, 372
Determining the Sample Size: Ad Hoc Methods, 373
Population Characteristics/Parameters, 374
Sample Characteristics/Statistics, 375
Sample Reliability, 376
Interval Estimation, 378
Sample Size Question, 380
Proportions, 381
Coefficient of Variation, 383
Stratified Sampling, 383
Multistage Design, 386
Sequential Sampling, 386
Summary, 387
Questions and Problems, 387
End Notes, 389
Cases for Part II, 390
Case II-1: Currency Concepts International, 390
Appendix A: Interview Guide for International Travelers (U.S. Citizens), 394

Part III Data Analysis

16 Fundamentals of Data Analysis 396

Learning Objectives, 396
Preparing the Data for Analysis, 397
Strategy for Data Analysis, 401
Cross-Tabulations, 405
Factors Influencing the Choice of Statistical Technique, 407
An Overview of Statistical Techniques, 409
Summary, 414
Questions and Problems, 414
End Notes, 415
Appendix: Choice of a Statistical Package, 415

17 Hypothesis Testing: Basic Concepts and Tests of

Associations 416
Learning Objectives, 416
The Logic of Hypothesis Testing, 417
Steps in Hypothesis Testing, 418
Basic Concepts of Hypothesis Testing, 419
Cross-Tabulation and Chi-Square, 422
 Contents xxi

Summary, 431
Questions and Problems, 431
End Notes, 432
Case 17-1: Medical Systems Associates: Measuring Patient Satisfaction, 433

18 Hypothesis Testing: Means and Proportions 435

Learning Objectives, 435
Commonly Used Hypothesis Tests in Marketing Research, 436
Relationship Between Confidence Interval and Hypothesis Testing, 439
Analysis of Variance (ANOVA), 440
The Probability‐Values (p‐Values) Approach to Hypothesis Testing, 454
Effect of Sample Size and Interpretation of Test Results, 455
Summary, 456
Questions and Problems, 456
End Notes, 457
Case 18-1: American Conservatory Theater, 458
Case 18-2: Apple Appliance Stores, 459

Part IV Special Topics in Data Analysis

19 Correlation Analysis and Regression Analysis 461

Learning Objectives, 461
Correlation Analysis, 462
Regression Analysis, 466
Summary, 485
Questions and Problems, 486
End Notes, 488
Case 19-1: The Seafood Grotto, 489
Case 19-2: Election Research, Inc., 490

20 Discriminant, Factor, and Cluster Analysis 491

Learning Objectives, 491
Discriminant Analysis, 491
Multiple Discriminant Analysis, 500
Summary of Discriminant Analysis 504
Factor and Cluster Analysis, 504
Factor Analysis, 505
Summary of Factor Analysis, 516
Cluster Analysis, 517
Summary of Cluster Analysis, 527
Questions and Problems, 527
End Notes, 529
Case 20-1: Southwest Utility, 529
Case 20-2: Store Image Study, 530
xxii Contents

21 Multidimensional Scaling and Conjoint Analysis 533

Learning Objectives, 533
Multidimensional Scaling, 533
Attribute‐Based Approaches, 534
Summary of MDS, 543
Conjoint Analysis, 544
Summary of Conjoint Analysis, 551
Questions and Problems, 552
End Notes, 553
Case 21-1: Nester’s Foods, 554
Case 21-2: The Electric Truck Case, 555

22 Presenting the Results 556

Learning Objectives, 556
Guidelines for Successful Presentations, 556
Written Report, 562
Oral Presentation, 566
Relationship with the Client, 569
Summary, 570
Questions and Problems, 570
End Notes, 570

Part V Applications of Marketing Intelligence

23 Marketing-Mix Measures 571

Learning Objectives, 571
New-Product Research 572
Pricing Research 582
Distribution Research, 585
Promotion Research, 591
Total Quality Management, 601
Summary, 606
Questions and Problems, 607
End Notes, 609
Case 23-1: National Chemical Corporation, 610
Case 23-2: Hokey Pokey is Born in India, 611

24 Brand and Customer Metrics 613

Learning Objectives, 613
Competitive Advantage, 613
Brand Equity, 616
Customer Satisfaction 622
Contemporary Applications of Marketing Intelligence 628
 Contents xxiii

Maximizing Customer Profitability, 630

Summary, 633
Questions and Problems, 633
End Notes, 633

25 New Age Strategies 635

Learning Objectives, 635
Database Marketing, 636
Relationship Marketing, 645
E-Commerce, 649
Mobile Marketing, 652
Social Marketing, 654
Experiential Marketing 658
Word-of-Mouth Marketing, 659
Internet of Things, 662
Bots and Artificial Intelligence, 664
Blockchain, 666
Summary, 668
Questions and Problems, 669
End Notes, 669

Appendix A 673
A-1. Standard Normal, Cumulative Probability in Right‐Hand Tail for
Positive Values of z; Areas are Formed by Symmetry 673
A-2. χ 2 Critical Points 674
A-3. F Critical Points 676
A-4. Cut‐Off Points for the Student’s t‐Distribution 681
A-5. Procedures for Conducting Univariate and Multivariate Analysis in SPSS 682
A-6. Output of Select Tables in SPSS 686

Glossary 699
Index 717
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 Fig. 144. Lepidodendron Veltheimianum Sternb.
A. Leaf-cushion and leaf-scar seen in surface-view at a; on the rest
of the specimen a slightly lower surface is exposed. (After Stur.)
B. Diagrammatic longitudinal section to explain the differences
between its two surfaces a and b shown in fig. A.
The shaded portion c represents the rock matrix, the surfaces
ab, ed, mark the outer and inner edge of the outer portion of the
bark of the Lepidodendron stem.
lt, leaf-trace; p, p′, parichnos.
 Fig. 145.
A. Diagrammatic surface-view and longitudinal section of a
Lepidodendron leaf-cushion.
B. Aerenchyma below the leaf-scar. (After F. E. Weiss.)

The parichnos scars are shown on the leaf-scar and cushion in fig.
146, C. In the lower leaf-cushion shown in fig. 146, E, the infra-foliar
parichnos scars, p, are clearly seen, but the preservation of the leaf-
scar is not sufficiently good to show them on that part of the fossil. In
the upper cushion (fig. 146, E) the position of the parichnos arms is
shown on the leaf-scar, but the infra-foliar parichnos scars are
hidden by two small spiral shells. The genus Spirorbis, to which
these shells are referred, appears to have persisted from the Silurian
epoch to the present day. The comparatively frequent occurrence of
Spirorbis shells on the leaves and other parts of Palaeozoic plants,
has recently been dealt with in a paper by Barrois[245] who discusses
in detail the habitats of these small animals from the point of view of
the conditions under which the plants were preserved. In a note by
Malaquin appended to Barrois’ paper the belief is expressed that
Spirorbis lived on pieces of Palaeozoic plants which lay under water.
The fact that with one exception all the Spirorbis shells on the
specimen of Lepidodendron, of which two leaf-cushions are shown in
fig. 146, E, occur on the large parichnos scars on the cheeks of the
cushions, suggests the possibility that the escape of gases from the
parichnos tissue may have rendered the position attractive to the
Spirorbis. It can hardly be accidental that the shells occur on the
parichnos strands. This fact recalls the view held by Binney[246] and
accepted with favour by Darwin[247] that Lepidodendron and other
coal-forest trees may have lived with the lower parts of the stems in
sea water.
Above the leaf-scar is a fairly deep triangular or crescentic pit (fig.
146, C, l) known as the ligular pit from the occurrence on younger
shoots of a delicate organ like the ligule of Isoetes (fig. 132)
embedded in a depression in the upper part of the leaf-cushion. The
ligule was first figured in Lepidodendron by Solms-Laubach[248] and
described in English material by Williamson under the name of the
adenoid organ[249].
In some Lepidodendron stems a second triangular depression
may occur above the ligular pit, the meaning of which is not clear:
this has been called the triangulum by Potonié[250]. Stur[251] suggested
that it may represent the position occupied by a sporangium in
Lepidodendron cones.
It is important to remember that as a branch increases in girth the
leaf-cushions are capable of only a certain amount of growth: when
the limit is reached they are stretched farther apart and thus the
narrow groove which separates them is converted in older stems into
a comparatively broad and flat channel, thus altering the surface
characters.
 Fig. 146. Lepidophloios and Lepidodendron leaf-cushions.
A, B, D, F, G, H, I. Lepidophloios. (Fig. A should be reversed.)
C, E. Lepidodendron aculeatum.
A, B. From a specimen in the Sedgwick Museum, Cambridge (leaf-
cushion 3 cm. broad).
C. From a specimen in the Sedgwick Museum, Cambridge (leaf-cushion 4
cm. long).
D. From a section in the Cambridge Botany School Collection.
E. From a specimen in the Bunbury Collection, Cambridge Botany
School, showing Spirorbis shells (leaf-cushion 2 cm. long).
F. From a section in the Williamson Collection, British Museum No. 1,
973.
G, H, I. From sections in the Cambridge Botany School Collection.

Another feature worthy of notice in reference to the leaf-cushions

of Lepidodendron is the occurrence in rare instances of alternate
zones of larger and smaller cushions. This variation in the size of the
leaf-cushions is by no means uncommon in the closely allied genus
Sigillaria; in Lepidodendron it has been described by Potonié[252] in L.
volkmannianum and more recently by Mr Leslie and myself[253] in a
South African species L. vereenigense.
Owing to the natural exfoliation of the superficial layers of the
outer bark at a certain stage in the growth of the plant, or in some
instances no doubt as the result of post-mortem decay, which
destroys the delicate cells of the meristematic zone in the outer
cortex, isolated leaf-cushions and strips of the external surface are
occasionally met with as carbonised impressions.
The appearance presented by a Lepidodendron stem which has
been deprived of its superficial tissues may be dealt with more
intelligibly after we have become familiar with the anatomical
characters.

iii. Lepidophloios.
Before proceeding further with the genus Lepidodendron a short
account may be intercalated of the external features of a
lepidodendroid type of stem which it is customary to describe under
a distinct generic title Lepidophloios. This name is convenient for
diagnostic purposes though it seems clear that apart from the form of
the leaf-cushion (fig. 146, A) we are at present unable to recognise
any well-defined differences between the two forms Lepidodendron
and Lepidophloios. For general purposes the name Lepidodendron
will be used as including plants possessing leaf-cushions of the type
already described as well as those with the Lepidophloios form of
cushion.
The generic name Lepidophloios was first used by Sternberg[254]
for a Carboniferous species which he had previously described as
Lepidodendron laricinum. In 1845 Corda[255] instituted the name
Lomatophloios for specimens possessing the same external
characters as those for which Sternberg had chosen the name
Lepidophloios. The leaf-cushions of Lepidophloios differ from those
of the true Lepidodendron in their relatively greater lateral extension
(cf. fig. 146, A and C), in their imbricate arrangement and in bearing
the leaf, or leaf-scar, at the summit. In some species referred to
Lepidophloios the cushions are however vertically elongated and in
this respect similar to those of Lepidodendron: an example of this
type is afforded by Lepidophloios Dessorti a French species
described by Zeiller[256]. In younger branches the cushions may be
directed upwards having the leaf-scar at the top; but in the majority
of specimens the cushions are deflexed as in figs. 146, D; 160, A.
The shoot of Lycopodium dichotomum shown in fig. 121, B, with the
leaves in the reversed position bears a close resemblance to a
branch of Lepidophloios.
The photograph of Lepidophloios scoticus Kidst.[257] reproduced in
fig. 160, A, illustrates the dichotomous branching of the stem and the
form of the cushions with the leaf-scars pointing downwards. In the
fertile branch of the same species shown in fig. 160, B, the leaf-scars
face upwards.
In most species the cushions are simply convex without a median
keel, but in some cases a median ridge divides the cushion into two
cheeks as in the genus Lepidodendron. The leaf-scar bears three
small scars, the larger median scar marking the position of the leaf-
trace, while the lateral scars are formed by the two arms of the
parichnos: in some examples of deflexed cushions, though not in all,
a ligular pit occurs on the cushion a short distance above the leaf-
scar.
The drawing reproduced in fig. 146, A, showing the leaf-scar on
the upper edge of the cushion should have been reversed with the
leaf-scars pointing downwards. This figure represents part of the
surface of a specimen consisting of the outer cortex of a stem with
leaf-cushions 3 cm. broad. The thickness of this specimen is 4 cm.: a
section through the line ab is represented in fig. 146, D (reproduced
in the correct position, with the leaf-scars, sc, pointing downwards):
internal to the cushions is a band of secondary cortex (the shaded
strip on the outer edge of the section) which was formed on the
outside of the phellogen. The phellogen is a cylinder of actively
dividing cells in the outer part of the cortex of the stem, often spoken
of as the cork-cambium or cortical meristem, which produces a
considerable amount of secondary cortical tissue on its inner face
and a much smaller amount towards the stem surface. This delicate
cylinder frequently forms a natural line of separation between the
outer shell of bark and the rest of the stem. In the specimen before
us, the thin-walled cells of the phellogen were ruptured before
petrification and the outer shell of bark was thus separated as a
hollow cylinder from the rest of the stem: this cylinder was then
flattened, the two inner surfaces coming into contact. Fig. 146, D,
represents a section of one half of the thickness of the flattened
shell.
This separation of the outer cortex, and its preservation apart from
the rest of the stem, is of frequent occurrence in fossil
lycopodiaceous stems. The flattened outer cortical shell of a
Lepidophloios, specifically identical with that shown in fig. 146, A and
D, was erroneously described by Dr C. E. Weiss in 1881 as a large
lepidodendroid cone[258].
Fig. 146, B, affords a view of the inner face of the specimen of
which the outer surface is seen in fig. 146, A: the surface shown in
the lower part of the drawing, on which the boundaries of the
cushions are represented by a reticulum, corresponds to the inner
edge of the strip of secondary cortical tissue represented by the
vertically shaded band in fig. 146, D.
The shaded surface in fig. 146, B, represents a slightly deeper
level in the stem which corresponds to the outer edge of the
vertically shaded band of fig. 146, D: the narrow tapered ridges (fig.
146, B) represent the leaf-traces passing through the secondary
cortex, and the fine vertical shading indicates the elongated
elements of which this strip of secondary cortex is composed.
In the longitudinal section diagrammatically reproduced in fig. 146,
D, cut along the line ab of fig. 146, A, the parenchymatous tissue of
the stout cushions has been partially destroyed, as at a; at s is seen
the section of a Stigmarian rootlet which has found its way into the
interior of a cushion. Each leaf-trace is accompanied by a parichnos
strand as in the true Lepidodendron; at the base of the leaf-cushion
the parichnos branches into two arms which diverge slightly right and
left of the leaf-trace, finally entering the base of the leaf lamina as
two lateral strands (fig. 147, p). At one point in fig. 146, D the section
has shaved a leaf-trace represented by a black patch resting on the
parichnos just above the line ef, but it passes through one of the
parichnos arms p′ which debouches on to the leaf-scar sc at p. Had
the section been cut along the line cd of fig. 146, A the leaf-trace
would have been seen in a position similar to that occupied by the
parichnos p′ in fig. 146, D.
 Fig. 147. Lepidophloios leaf-cushion in tangential section. (From a
section in the Williamson Collection, British Museum, No. 1973.)
Fig. 147, A, affords a good example of a tangential section through
a Lepidophloios leaf-cushion, 1 cm. broad, like that represented in
fig. 146, A, showing the vascular bundle lt, the two parichnos
strands, p, composed of large thin-walled cells (cf. Isoetes, fig. 133,
H, I), and the ligular pit near the upper edge of the section enclosing
the shrunken remains of the ligule (fig. 147, B, l).
LEPIDODENDRON

Fig. 147, B, shows the form of the tangentially elongated leaf-

cushions of Lepidophloios and their spiral disposition.
 Fig. 146, F, represents a section similar to that shown in figs. 147,
A and B, but in this case the leaf-trace, lt, and the parichnos strands,
p, lie in a cavity formed by the destruction of some of the leaf-
cushion tissue. It is worthy of notice that the parichnos cells have
resisted decay more successfully than the adjacent tissue of the
cushion.
The diagrammatic sketches reproduced in fig. 146, H and I, were
made from a transverse section similar to one originally figured by
Williamson[259]: fig. 146, H, corresponding in position to the line gh in
fig. 146, A, passes through the ligular pit, l, and cuts across the
parichnos in the act of branching; the leaf-trace passes outwards
beyond the Y-shaped parichnos strand. In the other section, fig. 146,
I, the parichnos is shown in a horizontal plane and the leaf-trace, lt,
appears in oblique transverse section. In both sections and in fig.
146, G the shaded band at the base represents the secondary
cortical tissue external to the phellogen.
The transverse section represented in fig. 146, G, shows in the
left-hand cushion, a, the exit of the two parichnos arms and the leaf-
trace between them: it illustrates also the various forms assumed by
lepidodendroid leaf-cushions when cut across at different levels.

iv. The Anatomy of Lepidodendron vasculare

Binney[260].
Figs. 148–155, 168, A.
In the earlier literature dealing with the anatomy of Lepidodendron
and Sigillaria the presence or absence of secondary vascular tissue
was made the criterion of generic distinction and the distinguishing
feature between the classes Pteridophytes and Gymnosperms,
Lepidodendron being relegated to the former class because it was
supposed to have no power of forming secondary wood, while
Sigillaria, characterised by a considerable development of such
tissue, was classed by Brongniart and afterwards by Renault as a
Gymnosperm. Binney[261] in 1865 recognised that the two types of
stem pass into one another, but it was Williamson[262] who provided
complete demonstration of the fallacy of the Brongniartian view.
These two undoubted Pteridophytes agree very closely in
anatomical structure and both are now recognised as arborescent
genera of Lycopodiaceous plants. In a paper published by Lomax
and Weiss in 1905[263] a specimen is described from the Coal-
Measures of Huddersfield, in which a decorticated stem with the
anatomical characters of Binney’s Sigillaria vascularis gives off a
branch having the anatomical structure which it has been customary
to associate with the species Lepidodendron selaginoides, so-called
by Sternberg and founded by him on impressions showing well-
preserved external characters.
In 1862 Binney[264] described petrified specimens of vegetative
shoots from the Lower Coal-Measures of Lancashire under the
names Sigillaria vascularis and Lepidodendron vasculare. These
were afterwards recognised as different states of the same species.
A few years after the publication of Binney’s paper Carruthers[265]
identified Binney’s species Lepidodendron vasculare with
Sternberg’s L. selaginoides. The evidence on which this identification
rests has not been stated, but many writers have retained this
specific designation for the well-defined type of anatomical structure
first described by Binney as L. vasculare. The use of the specific
name selaginoides is, however, open to objection. The species
Lepidodendron selaginoides, as pointed out by Kidston[266], is
probably identical with the plant which Brongniart had named L.
Sternbergii before the institution of Sternberg’s species, and we are
not in possession of convincing evidence as to the connection of L.
Sternbergii (= L. selaginoides) with specimens possessing the
anatomy of Binney’s type. Binney’s designation is therefore retained
for the anatomical type described in the following pages[267].
The most detailed account hitherto published of the anatomy of
Lepidodendron vasculare is that by the late M. Hovelacque[268],
based on material from the Lower Coal-Measures of England.
 Fig. 148. Lepidodendron vasculare Binney.
A. Transverse section. (Based on a section 2·5 cm. in diameter, in
the Cambridge Botany School Collection.)
B. Longitudinal section. (Drawn from a section in Dr Kidston’s
Collection.)

The small shoot, represented somewhat diagrammatically in fig.

148, A, illustrates the anatomical features of a typical example of the
species: the shoot has a diameter of 2·5 cm. and its central cylinder
(x-sc) is 2·5 mm. in width.
 Noticeable features are (i) the small size of the central cylinder (or
stele) in proportion to the diameter of the branch, (ii) the production
at a comparatively early stage of growth of a zone of secondary
wood, x2, which gradually assumes the form of a complete cylinder
of unequal breadth, surrounding the primary xylem, x, (iii) the
formation of a secondary cortical tissue by a meristematic cylinder
(phellogen, pl) situated close to the leaf-cushion region of the outer
cortex. On the outer edge the stele consists of narrow tracheae
some of which show in longitudinal section the spiral form of
thickening characteristic of most protoxylem elements: towards the
centre of the stele the diameter of the tracheae gradually increases
and parenchymatous cells become associated with the elongated
scalariform elements. In the central region the stele is composed of
parenchymatous tissue arranged in vertical series of short cells,
interspersed with short tracheae distinguished by the greater
thickness of their walls and by their scalariform and reticulate
thickening bands. Some of these short tracheae are shown in
vertical section in fig. 149, B: the fine and broken lines connecting
adjacent thickening bands probably represent the remains of the
original wall. These delicate bands, which have been figured in
various species of lepidodendroid plants[269], are worthy of notice in
connexion with the recent work of Mr Gwynne-Vaughan[270] who has
shown that in many recent ferns the scalariform bands in the xylem
elements are not connected by a thin pit-closing membrane, but are
separated from one another by open spaces. In the Lepidodendron
tracheae we seem to have a stage in which the intervening
membrane is in process of absorption. It is, however, possible that
the threads may be the result of contraction and splitting of the
membrane during drying or decay.
 Fig. 149. Lepidodendron vasculare. a, immature tracheae; m, meristem;
mr, medullary ray; x, xylem.
A. Longitudinal section through the edge of the secondary wood.
B. Short tracheae in the centre of the stele. (From a specimen from
the Halifax Hard bed in Dr Kidston’s Collection.)

The stele of Lepidodendron vasculare, before the addition of any

secondary xylem, may be described as a protostele, a term originally
proposed by Professor Jeffrey[271], in which the central part of the
conducting strand of xylem elements has been converted into rows
of parenchyma and short tracheids, the latter being better adapted to
storage than to conduction. It is probable that this type of stelar
anatomy, which distinguishes L. vasculare from other species,
represents a comparatively primitive arrangement forming a
transition between the stele of L. esnostense, which consists of a
solid rod of tracheids, and the stele of L. Harcourtii (fig. 179, A) and
other species in which the xylem forms a cylinder enclosing a large
parenchymatous pith.
Parenchymatous cells occur in contact with the outer edge of the
xylem-cylinder some of which are distinguished by an irregular
reticulate pitting. The tangential section represented in fig. 148, B,
illustrates the appearance of a shoot of L. vasculare in which no
secondary xylem is present: the central strand of tissue consists of
the parenchyma abutting on the xylem with several leaf-traces (lt)
passing upwards in an almost vertical course from the outer edge of
the stele.
The secondary xylem (fig. 148, A, x2) consists of radially arranged
scalariform tracheae with associated rows of parenchymatous cells
which form medullary rays (fig. 149, mr). Leaf-traces pass through
the medullary rays in the secondary xylem cylinder in a direction at
right angles to the primary xylem stele from which they are given off,
but at the outer edge of the secondary xylem they bend suddenly
upwards and for a time follow a steep and almost vertical course.
In well-preserved longitudinal sections the outermost secondary
xylem tracheae are seen to be succeeded by a few narrow and
vertically elongated elements (fig. 149, A, a), which represent young
unlignified tracheae: these are followed by shorter parenchymatous
cells (m) forming part of a meristematic zone from which the
secondary xylem receives additions.
Returning to fig. 148, A; the zone of secondary wood, x2,
composed of scalariform tracheids and medullary rays, is succeeded
by a few layers of parenchymatous cells and beyond this is a
broader zone, sc, to which the term secretory zone has been
applied[272]; this is made up of small parenchymatous cells varying in
size and of larger spaces which appear to have been formed by the
disorganisation of thin-walled elements. The whole zone presents a
characteristic appearance due to the association of small cells, large
clear spaces, and a certain amount of dark-coloured material
suggestive of tissue disorganisation and secreted products. The
anatomical characters of the secretory zone are shown in the
photograph, fig. 168, A, sc. Several leaf-traces are seen in
transverse section in the secretory zone (black dots in fig. 148, A, sc;
fig. 154, C, lt): each trace consists of a strand of narrow tracheae
accompanied by a few encircling layers of small parenchymatous
cells. As a trace continues its steeply ascending course through the
secretory zone, it becomes associated with a strand of that tissue
and assumes the form of a collateral vascular bundle, the outer part
of which does not consist of typical phloem but of shorter elements
derived from the secretory zone. Beyond the secretory zone we find
a more homogeneous tissue composed of parenchymatous
elements slightly extended tangentially (figs. 148, A, c1; fig. 168, A,
c); this is spoken of as the inner cortical region. In the great majority
of sections of L. vasculare as of other species of the genus, the
broader middle cortex (fig. 148, c2) is occupied by mineral matter,
introduced subsequent to decay of the tissue; or it is represented by
patches of delicate tissue composed of loosely arranged
parenchymatous cells varying considerably in size and shape, some
being small, oval or polygonal elements while others have the form
of sinuous hypha-like tubes.
In this middle cortical region may be seen leaf-traces passing
outwards in an almost horizontal course (fig. 148, A, lt): after leaving
the inner cortex the leaf-traces bend somewhat abruptly outwards to
follow a more direct path through the middle and outer cortex. The
ring of tissue, s, seen in the middle cortex of fig. 148, A, belongs to a
Stigmarian rootlet.
The outer cortex (fig. 148, A and B, c3) consists of homogeneous
parenchyma which is stronger and more resistant to decay than the
looser middle cortex. The leaf-traces, as shown in fig. 148, B, pass
through this region in a rather steeply ascending direction: each is
seen to be enclosed by a space originally occupied by a strand of
middle cortical tissue which accompanies lepidodendroid leaf-traces
on their under side and has already been described as the
parichnos, (pp. 97, 100–103; figs. 146, 147).
The surface of the stem shown in section in fig. 148, A, is
composed of broad leaf-cushions. A single leaf-trace with its
parichnos passes into each cushion, but in the neighbourhood of the
base of a cushion the parichnos bifurcates (cf. fig. 146, H, I) and the
arms diverge slightly to the right and left finally passing beyond the
cushion into the lamina of the leaf, their position being shown, as
already explained, by the two small lateral scars on the leaf-scar
area.
The diagrammatic sketch of a radial longitudinal section through a
leaf-cushion represented in fig. 150 illustrates the relation of the leaf-
trace to the leaf-cushion. The trace consists of xylem, x, above and a
strand of the secretory zone, st, below; the parichnos tissue was
originally present on the under side of the leaf-trace at a. The
external surface, bc, marks the limit of the leaf-scar through the
middle of which passes the vascular strand lt.
The lower gap a has been formed by the tearing of thin-walled
cells of the phellogen, the meristematic tissue from which a
considerable amount of secondary cortical tissue or phelloderm has
been produced at pd. On the outside of the cushion, c, the cells are
somewhat crushed and distinguished by their darker colour from the
bulk of the parenchymatous tissue d.
This section also illustrates another characteristic feature of
Lepidodendron, namely the presence of a ligule and a ligular pit: the
former is represented by a carbonised patch of tissue and the latter
extends from the surface of the cushion at b, just above the leaf-
scar, almost to the level of the leaf-trace, lt. A comparison of this
section with figs. 146 and 147 will make clear the relation of the
several parts of the cushion and leaf-scar.
The gaps gg, seen in fig. 148, A and B, mark the position of the
delicate meristematic zone or phellogen which arises close to the
bases of the leaf-cushions; the phellogen has already produced a
few rows of radially disposed elements, represented by short radial
lines in the drawing, which constitute secondary cortical tissue.
 Fig. 150. Lepidodendron vasculare. Leaf-cushions in longitudinal section.
(From a specimen in Dr Kidston’s Collection.)
In older shoots the amount of the secondary cortical tissue
developed on the inner side of the phellogen is considerable (cf. figs.
152, 153).
The structure of the cortex of a shoot in which secondary growth,
both in the stele and in the outer cortex, has progressed further than
in the specimen shown in fig. 148 is represented in fig. 151.
 Fig. 151. Lepidodendron vasculare. An older stem than that shown in fig.
148. (From a section in the Manchester Museum. No. 351.)
The section (fig. 151, A) measures 7 × 3·8 cm. in diameter; the
primary xylem is surrounded by a fairly broad cylinder of secondary
wood (fig. 151, E, x and x2). The almost smooth surface of the
primary wood (fig. 151, E, x) is succeeded by the secondary xylem,
x2, characterised at its inner edge by the tapered ends of the radial
rows of scalariform tracheids between which occur several delicate
parenchymatous cells (fig. 151, E, a). The occurrence of such
isodiametric elements, often exhibiting a delicate spiral thickening
band, is a characteristic feature of the boundary between primary
and secondary wood in lepidodendroid stems. The secondary wood
is penetrated by numerous medullary rays and in some of them are
seen strands of narrow spirally thickened tracheae—the leaf-traces
—which are in organic continuity with the exarch protoxylem of the
primary wood. The leaf-traces are oval and mesarch. The space, c2,
(fig. 151, A) originally occupied by the delicate middle cortex, is
succeeded by a shell of outer cortex composed chiefly of secondary
tissue (phelloderm, pd) passing towards the inner boundary of this
region into the primary outer cortex g (fig. 151, A and C). The radially
disposed elements which make up the bulk of the phelloderm are
associated with concentric rows of secretory strands, represented by
tangentially arranged dots in fig. 151, A: on the outer edge of the
phelloderm a few patches of primary cortex are still preserved, as at
c, fig. A. One of these is shown on a larger scale in fig. B; at m the
phelloderm is interrupted by a gap beyond which the cells have
thinner walls and show signs of recent division; this is probably the
position of the phellogen. The tissue b, fig. 151, B, consists of
secondary cortex succeeded beyond d by the parenchymatous
tissue of the leaf-cushion, in which the remains of a ligule, l, are seen
in the ligular pit. This section corresponds in position to a line drawn
across fig. 150 at the level of b. In this specimen we have two kinds
of secondary cortical tissue: that formed external to the phellogen,
from m to d in fig. 151, B, is less in amount than that produced
internal to the phellogen. We cannot make any satisfactory
statement as to the nature of this secondary tissue, whether or not
any of it agreed in composition with the cork which is usually formed
external to the phellogen in recent plants. As the stem of a
Lepidodendron grew in girth the leaf-cushions became separated by
intervening depressions composed of the secondary cortex formed
external to the phellogen, but at a later stage the cushions were
thrown off, leaving the outer edge of the phelloderm as the
superficial tissue. This exposed tissue became fissured as growth
and consequent stretching continued, producing the appearance
seen on the surface of the still older stem represented in fig. 153.
The inner edge of the phelloderm seen at e in fig. 151, C, passes
suddenly into the inner primary region of the outer cortex (fig. 151, A
and C, g) which comprises two types of parenchymatous tissue,
patches of isodiametric cells, g, g, alternating with radially arranged
areas consisting of tangentially elongated elements (fig. C, f, f; fig. D)
which extend as wedges into the phelloderm.
The longitudinal section represented in fig. 152, B, shows an equal
bifurcation of a stem in which no secondary xylem is present; in the
lower part of the section the xylem and the outgoing leaf-traces are
seen in radial section and at the upper end of each arm the leaf-
traces alone, lt, are exposed, as in fig. 148, B. It is interesting to