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Table of Contents

Cross – database joins 75

Blending data sources 76
Blending example 79
Filtering data 80
Filtering discrete fields 81
Filtering continuous fields 83
Filtering dates 85
Other filtering options 86
Summary 87
Chapter 3: Moving from Foundational to More Advanced Visualizations 88
Comparing values across different dimensions 89
Bar charts 89
Bar chart variations 92
Bullet chart – showing progress toward a goal 92
Bar in bar chart 95
Highlighting categories of interest 98
Visualizing dates and times 100
The built-in date hierarchy 101
Variations of date and time visualizations 105
Gantt charts 107
Relating parts of the data to the whole 110
Stacked bars 111
Treemaps 114
Area charts 117
Pie charts 119
Visualizing distributions 120
Circle charts 121
Jittering 122
Box and whisker plots 124
Histograms 125
Visualizing multiple axes to compare different measures 129
Scatterplot 129
Dual axis 131
Combination charts 133
Summary 134
Chapter 4: Using Row-Level, Aggregate, and Level of Detail
Calculations 135
Creating and editing calculations 137
Overview of the three main types of calculations 141
Row Level examples 142
Aggregate Level example 145
Row Level or Aggregate – why does it matter? 147
Level of Detail calculations 150
Level of Detail syntax 150

[ ii ]
Table of Contents

Level of Detail example 152

Parameters 156
Creating parameters 156
Practical examples of calculations and parameters 160
Fixing data issues 160
Extending the data 161
Enhancing user experience, analysis, and visualizations 163
Achieving flexibility with data blends 166
Ad hoc calculations 172
Performance considerations 175
Summary 176
Chapter 5: Table Calculations 177
Overview of table calculations 178
Creating and editing table calculations 179
Quick table calculations 181
Relative versus fixed 183
Scope and direction 184
Working with scope and direction 186
Addressing and partitioning 189
Advanced addressing and partitioning 192
Custom table calculations 194
Practical examples 199
Year – over – Year growth 199
Ranking within higher levels 201
Late filtering 203
Data densification 204
When and where data densification occurs 205
An example of leveraging data densification 209
Summary 215
Chapter 6: Formatting a Visualization to Look Great and Work Well 216
Formatting considerations 216
Understanding how formatting works in Tableau 218
Worksheet level formatting 219
Field-level formatting 224
Additional formatting options 230
Adding value to visualizations 232
Tooltips 237
Summary 238
Chapter 7: Telling a Data Story with Dashboards 239
Dashboard objectives 240
Example - is least profitable always unprofitable? 242
Building the views 242

[ iii ]
Table of Contents

Creating the dashboard framework 245

Implementing actions to tell the story 250
Designing for different displays and devices 255
How actions work 259
Filter actions 260
Highlight actions 263
URL actions 265
Example - regional scorecard 265
Stories 270
Summary 278
Chapter 8: Deeper Analysis - Trends, Clustering, Distributions, and
Forecasting 279
Trending 280
Customizing trend lines 285
Trend models 289
Analyzing trend models 293
Clustering 299
Distributions 304
Forecasting 308
Summary 314
Chapter 9: Making Data Work for You 315
Structuring data for Tableau 315
Good structure - tall and narrow instead of short and wide 317
Wide data 317
Tall data 318
Wide and tall in Tableau 319
Good structure - star schemas 322
Techniques for dealing with data structure issues 325
Restructuring data in Tableau connections 326
Union files together 331
Originals 332
Prequels 332
Sequels 332
Cross-database joins 336
Working with different Level of Detail 341
Overview of advanced fixes for data problems 346
Summary 348
Chapter 10: Advanced Visualizations, Techniques, Tips, and Tricks 349
Advanced visualizations 350
Slope chart 350
Lollipop chart 352
Waterfall chart 354
Sparklines 355

[ iv ]
Table of Contents

Dumbbell chart 356

Unit chart/symbol chart 358
Marimekko chart 361
Sheet swapping and dynamic dashboards 363
Dynamically showing and hiding other controls 369
Advanced mapping techniques 372
Supplementing the standard in geographic data 372
Manually assigning geographic locations 373
Creating custom territories 375
Ad hoc custom territories 376
Field – defined custom territories 378
Some final map tips 380
Using background images 382
Animation 386
Summary 387
Chapter 11: Sharing Your Data Story 388
Presenting, printing, and exporting 388
Presenting 389
Printing 389
Exporting 392
Sharing with users of Tableau Desktop and Tableau Reader 393
Sharing with Tableau Desktop users 393
Sharing with Tableau Reader users 394
Sharing with users of Tableau Server, Tableau Online, and Tableau
Public 395
Publishing to Tableau Public 396
Publishing to Tableau Server and Tableau Online 397
Interacting with Tableau Server 400
Additional distribution options using Tableau Server 402
Summary 402
Chapter 12: Catching Up with Tableau 2018 403
Tableau Desktop 404
Data Source improvements 404
Normalized extract (2018.3) 404
Spatial join (2018.2) 406
Other Data Source improvements 410
Visualization improvements 410
Density Mark (2018.3) 410
Step and jump lines (2018.1) 414
Worksheet transparency (2018.3) 414
Dual Axis mapping (2018.1) 416
Nested sort (2018.2) 418
Hierarchy filtering (2018.1) 419
Other improvements 420
Dashboard improvements 421

[v]
Table of Contents

Extensions (2018.2) 421

Dashboard navigation button (2018.3) 423
Navigation action (2018.3) 426
The Change Set Values action (2018.3) 427
Automatic Mobile layouts (2018.2) 431
Grids (2018.2) 434
Tableau Server/Online 435
Interacting 435
Mixed content (2018.3) 435
Mobile preview (2018.3) 438
Comments (2018.2) 440
Web authoring 440
Connecting to data (2018.1) 440
Other web authoring improvements 441
Administration 441
Tableau Service Manager (2018.2) 442
Other administrative improvements 443
Summary 443
Chapter 13: Deal with Security 445
Tableau Server security 445
User Filters 448
Row-level filters 454
Summary 458
Chapter 14: How to Keep Growing Your Skills 459
The Tableau Community 459
Tableau Public 460
Community projects 462
Ambassadors, Zen Masters, and Iron Viz 464
Ambassadors 464
Zen Masters 464
Iron Viz 465
Summary 465
Other Books You May Enjoy 466
Index 469

[ vi ]
 Preface
Graphical presentation of data enables us to easily understand complex data sets. Tableau
10 Complete Reference provides easy-to-follow recipes with several use cases and real-
world business scenarios to get you up and running with Tableau 10.

This Learning Path begins with the history of data visualization and its importance in
today's businesses. You'll also be introduced to Tableau - how to connect, clean, and
analyze data in this visual analytics software. Then, you'll learn how to apply what you've
learned by creating some simple calculations in Tableau and using Table Calculations to
help drive greater analysis from your data. Next, you'll explore different advanced chart
types in Tableau. These chart types require you to have some understanding of the Tableau
interface and understand basic calculations. You’ll study in detail all dashboard techniques
and best practices. A number of recipes specifically for geospatial visualization, analytics,
and data preparation are also covered. Last but not least, you'll learn about the power of
storytelling through the creation of interactive dashboards in Tableau.

Through this Learning Path, you will gain confidence and competence to analyze and
communicate data and insights more efficiently and effectively by creating compelling
interactive charts, dashboards, and stories in Tableau.

Who This Book Is For

Tableau 10 Complete Reference is designed for anyone who wants to understand their data
better and represent it in an effective manner. It is also used for BI professionals and data
analysts who want to do better at their jobs.

What This Book Covers

Chapter 1, Creating Your First Visualizations and Dashboard, introduces the basic concepts of
data visualization and shows multiple examples of individual visualizations that are
ultimately put together in an interactive dashboard.

Chapter 2, Working with Data in Tableau, explains that Tableau has a very distinctive
paradigm for working with data. This chapter explores that paradigm and gives examples
of connecting to and working with various data sources.
Preface

Chapter 3, Moving from Foundational to More Advanced Visualizations, expands upon the
basic concepts of data visualization to show how to extend standard visualization types.

Chapter 4, Using Row-Level, Aggregate, and Level of Detail Calculations, introduces the
concepts of calculated fields and the practical use of calculations. The chapter walks
through the foundational concepts for creating Row Level, Aggregate, and Level of Detail
calculations.

Chapter 5, Table Calculations, is about table calculations, one of the most complex and most
powerful features of Tableau. This chapter breaks down the basics of scope, direction,
partitioning, and addressing to help you understand and use them to solve practical
problems.

Chapter 6, Formatting a Visualization to Look Great and Work Well, is about formatting, which
can make a standard visualization look great, have appeal, and communicate well. This
chapter introduces and explains the concepts around formatting in Tableau.

Chapter 7, Telling a Data Story with Dashboards, dives into the details of building
dashboards and telling stories with data. It covers the types of dashboards, objectives of
dashboards, and concepts such as actions and filters. All of this is done in the context of
practical examples.

Chapter 8, Deeper Analysis – Trends, Clustering, Distributions and Forecasting, explores the
analytical capabilities of Tableau and demonstrates how to use trend lines, clustering,
distributions, and forecasting to dive deeper into the analysis of your data.

Chapter 9, Making Data Work for You, shows that data in the real world isn’t always
structured well. This chapter examines the structures that work best and the techniques that
can be used to address data that can’t be fixed.

Chapter 10, Advanced Visualizations, Techniques, Tips, and Tricks, builds upon the concepts in
previous chapters and expands your horizons by introducing non-standard visualization
types along with numerous advanced techniques while giving practical advice and tips.

Chapter 11, Sharing Your Data Story, once you’ve built your visualizations and dashboards,
you’ll want to share them. This chapter explores numerous ways of sharing your stories
with others.

Chapter 12, Catching Up with Tableau 2018, details of every new feature of the different
Tableau 2018 versions. You'll learn how to use them with clear explanations, examples, and
tutorials. This chapter is the best way to catch up with the new releases if you already have
some Tableau knowledge.

[2]
Preface

Chapter 13, Deal with Security, is the last technical chapter of this book and focuses on three
ways to secure your data: permissions on Tableau Server, user filters on Tableau Desktop,
and row-level data security in your data.

Chapter 14, How to Keep Growing Your Skills, is a non-technical but essential chapter. You'll
discover many ways of learning new things and growing your Tableau skills thanks to
community projects. The chapter is also a tribute to the Tableau community, presenting
many ways to be part of that big family, which shares a passion for data visualization with
Tableau.

To Get the Most out of This Book

You will need a licensed or trial version of Tableau Desktop to follow the examples
contained in this book. You may download Tableau Desktop from Tableau Software
at http:/​/​www.​tableau.​com/​. The examples in this book use the interface and features of
Tableau 10.0. Many of the concepts will apply to previous versions, though some interface
steps and terminology may vary. The provided workbooks may be opened in Tableau 10.0
or later, though you may use any version to connect to the provided data files to work
through the examples. Tableau Public is also available as a free download (http:/​/​www.
tableau.​com/​) and may be used with many of the examples.

You may use a PC or a Mac to work through the examples in this book. Mac users may
notice slight changes in user interface and will need to make note of the following changes
in keys and clicks:

Right-click can be accomplished by holding the Command key while clicking

Right-click and drag and drop can be accomplished by holding the
option (Alt) key while dragging and dropping

Download the Example Code Files

You can download the example code files for this book from your account at
www.packt.com. If you purchased this book elsewhere, you can visit
www.packt.com/support and register to have the files emailed directly to you.

[3]
Preface

You can download the code files by following these steps:

1. Log in or register at www.packt.com.

2. Select the SUPPORT tab.
3. Click on Code Downloads & Errata.
4. Enter the name of the book in the Search box and follow the onscreen
instructions.

Once the file is downloaded, please make sure that you unzip or extract the folder using the
latest version of:

WinRAR/7-Zip for Windows

Zipeg/iZip/UnRarX for Mac
7-Zip/PeaZip for Linux

The code bundle for the book is also hosted on GitHub at https:/​/​github.​com/
PacktPublishing/​Tableau-​10-​Complete-​Reference. In case there's an update to the code,
it will be updated on the existing GitHub repository.

We also have other code bundles from our rich catalog of books and videos available
at https:/​/​github.​com/​PacktPublishing/​. Check them out!

Conventions Used
In this book, you will find a number of text styles that distinguish between different kinds
of information. Here are some examples of these styles and an explanation of their
meaning.

Code words in text, database table names, folder names, filenames, file extensions,
pathnames, dummy URLs, and user input are shown as follows: "We’ll create a calculated
field named Floor to determine if an apartment is upstairs or downstairs."

A block of code is set as follows:

IF [Apartment] >= 1 AND [Apartment] <= 3
THEN "Downstairs"
ELSEIF [Apartment] > 3 AND [Apartment] <= 6
THEN "Upstairs"
ELSE "Unknown"
END

[4]
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Preface

Bold: Indicates a new term, an important word, or words that you see onscreen. For
example, words in menus or dialog boxes appear in the text like this. Here is an example:
"When you open Tableau, on the left, in the Connect area, click on Microsoft Excel."

Warnings or important notes appear like this.

Tips and tricks appear like this.

Get in Touch
Feedback from our readers is always welcome.

General feedback: If you have questions about any aspect of this book, mention the book
title in the subject of your message and email us at [email protected].

Errata: Although we have taken every care to ensure the accuracy of our content, mistakes
do happen. If you have found a mistake in this book, we would be grateful if you would
report this to us. Please visit www.packt.com/submit-errata, selecting your book, clicking
on the Errata Submission Form link, and entering the details.

Piracy: If you come across any illegal copies of our works in any form on the Internet, we
would be grateful if you would provide us with the location address or website name.
Please contact us at [email protected] with a link to the material.

If you are interested in becoming an author: If there is a topic that you have expertise in
and you are interested in either writing or contributing to a book, please visit
authors.packtpub.com.

Reviews
Please leave a review. Once you have read and used this book, why not leave a review on
the site that you purchased it from? Potential readers can then see and use your unbiased
opinion to make purchase decisions, we at Packt can understand what you think about our
products, and our authors can see your feedback on their book. Thank you!

For more information about Packt, please visit packt.com.

[5]
 1
Creating Your First
Visualizations and Dashboard
Tableau is an amazing platform for seeing, understanding, and making key decisions based
on your data. With it, you can achieve incredible data discovery, analysis, and storytelling.
You'll accomplish these tasks and goals visually using an interface that is designed for a
natural and seamless flow of thought and work. Tableau accomplishes this using VizQL, a
visual query language. You won't have to learn VizQL. It's all done behind the scenes and
you won't be forced to write tedious SQL scripts, MDX code, or painstakingly work
through numerous wizards to select a chart type and then link everything to data.

Instead, you will be interacting with your data in a visual environment where everything
that you drag and drop will be translated into the necessary queries and then displayed
visually. You'll be working in real-time, so you will see results immediately, get answers as
fast as you can ask questions, and be able to iterate through dozens of ways to visualize the
data to find a key insight or tell a piece of the story.

Tableau allows you to accomplish numerous tasks, including:

Data connection, integration, and preparation: Tableau allows you to connect to

data from sources and, if necessary, create a structure that is ready to use. Most
of the time this is as easy as pointing Tableau to a database or opening a file, but
Tableau gives you the tools to bring together even complex and messy data from
multiple sources.
Data exploration: You can visually explore a dataset using Tableau in order to
understand what data you have.
Data visualization: This is the heart of Tableau. You can iterate through the
countless ways of visualizing the data to ask and answer questions, raise new
questions, and gain new insights.
Data analysis: Tableau has an ever growing set of analytical functions that allow
you to dive deep into understanding complex relationships, patterns, and
correlations in the data.
Creating Your First Visualizations and Dashboard Chapter 1

Data storytelling: Tableau allows you to build fully interactive dashboards and
stories with your visualizations and insights so that you can share the data story
with others.

We'll take a look at each of these tasks in the subsequent chapters. This chapter introduces
the foundational principals of Tableau and focuses on data visualization. We'll accomplish
this through a series of examples that will introduce the basics of connecting to data,
exploring and analyzing the data visually, and finally putting it all together in a fully
interactive dashboard. These concepts will be developed far more extensively in the
subsequent chapters. But don't skip this chapter, as it introduces key terminology and
foundational concepts, including:

Connecting to data
Foundations for building visualization
Visualizing the data
Creating bar charts
Creating line charts
Creating geographic visualizations
Using Show Me
Bringing everything together in a dashboard

Connecting to data
Tableau connects to data stored in a wide variety of files and databases. This includes flat
files, such as Excel and text files; relational databases, such as SQL Server and Oracle;
cloud-based data sources, such as Google Analytics and Amazon Redshift; and OLAP data
sources, such as Microsoft Analysis Services. With very few exceptions, the process of
building visualizations and performing analysis will be the same no matter what data
source you use. We'll cover the details of connecting to different data sources in Chapter 2,
Working with Data in Tableau.

For now, we'll connect to a text file, specifically, a comma-separated values file (.csv). The
data itself is a variation of the sample data provided with Tableau for Superstore, a fictional
retail chain that sells various products to customers across the United States. It's preferable
to use the supplied data file instead of the Tableau sample data as the variations will lead to
differences in visualizations.

[7]
Creating Your First Visualizations and Dashboard Chapter 1

The Chapter 1 workbook, included with the code files bundle, already have connections to
the file; however, for this example, we'll walk through the steps of creating a connection in
a new workbook:

1. Open Tableau; you should be able to see the home screen with a list of
connection options on the left, thumbnail previews of recently edited workbooks
in the center, links to various resources on the right, and sample workbooks on
the bottom.
2. Under Connect and To a file, click Text File.
3. In the Open dialogue box, navigate to the \Learning Tableau\Chapter 01\
directory and select the Superstore.csv file.
4. You will now see the data connection screen, which allows you to visually create
connections to data sources. We'll examine the features of this screen in detail in
the Connecting to data section of Chapter 2, Working with Data in Tableau. For now,
notice that Tableau has already added and given a preview of the file for the
connection:

[8]
Creating Your First Visualizations and Dashboard Chapter 1

5. For this connection, no other configuration is required, so simply click on the

Sheet 1 tab at the bottom to start visualizing the data! You should now see the
main work area within Tableau, which looks similar to the following screenshot:

We'll refer to elements of the interface throughout the book using specific terminology, so
take a moment to get familiar with the terms used for various components numbered in the
preceding image:

1. The menu contains various menu items for performing a wide range of functions.
2. The toolbar allows for common functions, such as undo, redo, save, adding a
data source, and so on.
3. The sidebar contains tabs for Data and Analytics. When the Data tab is active,
we'll refer to the sidebar as the data pane. When the Analytics tab is active, we'll
refer to the sidebar as the analytics pane. We'll go into detail later in this chapter,
but for now, note that the data pane shows the data source at the top and
contains a list of fields from the data source and is divided into dimensions and
measures.

[9]
Creating Your First Visualizations and Dashboard Chapter 1

4. Various shelves, such as Columns, Rows, Pages, and Filters, serve as areas to
drag and drop fields from the data pane. The Marks card contains additional
shelves, such as Color, Size, Text, Detail, and Tooltip. Tableau will visualize
data based on the fields you drop on the shelves.

Data fields in the data pane are available to be added to the view. Fields
that have been dropped on a shelf are called in the view or active fields,
because they play an active role in the way Tableau draws the
visualization.

5. The canvas or view is where Tableau will draw the data visualization. You may
also drop fields directly onto the view. In Tableau 10, you'll observe the seamless
title at the top of the canvas. By default, it will display the name of the sheet, but
it can be either edited or hidden.
6. Show Me is a feature that allows you to quickly iterate through various types of
visualizations based on data fields of interest. We'll look at Show Me towards the
end of the chapter.
7. The tabs at the bottom of the window gives you the option of editing the data
source, as well as navigating between and adding any number of sheets,
dashboards, or stories. Many times a tab (whether it is a sheet, dashboard, or
story) is referred to, generally, as a sheet. We'll also often use these specific terms
for a tab:
A sheet: A sheet is a single data visualization (such as a bar chart or
line graph). Since sheet is also a generic term for any tab, we'll often
refer to a sheet as a view because it is a single view of the data.
A dashboard: A dashboard is a presentation of any number of related
views and other elements (such as text or images) arranged together as
a cohesive whole to communicate a message to an audience.
Dashboards are often interactive.
A story: A story is a collection of dashboards or single views arranged
to communicate a narrative from the data. Stories can also be
interactive.

A Tableau workbook is the collection of data sources, sheets, dashboards,

and stories. All of this is saved as a single Tableau workbook file (.twb
or.twbx). We'll look at the difference in file types and explore details of
what else is saved as a part of a workbook in later chapters.

[ 10 ]
Creating Your First Visualizations and Dashboard Chapter 1

8. As you work, the status bar will display important information and details about
the view and selections.
9. Various controls allow you to navigate between sheets, dashboards, and stories,
as well as view the tabs as a filmstrip or switch to a Sheet Sorter showing an
interactive thumbnail of all sheets in the workbook.

Now that you have worked through connecting to the data, we'll explore some examples
that lay the foundation for data visualization and then move into building some
foundational visualization types. To prepare for this, do the following:

1. From the menu, navigate to File | Exit.

2. When prompted to save changes, select No.
3. From the \Learning Tableau\Chapter 01 directory, open the file Chapter
01 Starter.twbx. This file contains a connection to the Superstore data file
and is designed to help you walk through the examples in this chapter.

The files for each chapter include a Starter workbook that allows you to
work through the examples given in this book. If at any time, you'd like to
see the completed examples, open the Complete workbook for the
chapter.

With a connection to the data, you are now ready to visualize and analyze the data. As you
start doing so, you will take on the role of an analyst at the retail chain. You'll ask questions
of the data, build visualizations to answer those questions, and ultimately design a
dashboard to share the results. Let's start by laying down some foundations to understand
how Tableau visualizes data.

Foundations for building visualizations

When you first connect to a data source, such as the Superstore file, Tableau will display
the data connection and the fields in the data pane on the left sidebar. Fields can be dragged
from the data pane onto the canvas area or onto various shelves, such as Rows, Columns,
Color, or Size. We'll see that placement of the fields will result in different encodings of the
data, based on the type of field.

[ 11 ]
Creating Your First Visualizations and Dashboard Chapter 1

Measures and dimensions

The fields from the data source are visible in the data pane and are divided into measures
and dimensions. The difference between measures and dimensions is a fundamental
concept to understand when using Tableau:

Measures: Measures are values that are aggregated. That is, they can be
summed, averaged, and counted, or have a minimum or maximum.
Dimensions: Dimensions are values that determine the level of detail at which
measures are aggregated. You can think of them as slicing the measures or
creating groups into which the measures fit. The combination of dimensions used
in the view defines the view's basic level of detail.

As an example (which you can view in the Chapter 01 Starter workbook on the
Measures and Dimensions sheet), consider a view created using the fields Region and
Sales from the Superstore connection, as shown here:

[ 12 ]
Creating Your First Visualizations and Dashboard Chapter 1

The Sales field is used as a measure in this view. Specifically, it is being aggregated as a
sum. When you use a field as a measure in the view, the type aggregation (such as SUM,
MIN, MAX, AVG) will be shown on the active field. In the preceding example, the active
field on Rows clearly indicates the sum aggregation of Sales: SUM(Sales).

The Region field is a dimension with one of four values for each record of data: Central,
East, South, or West. When the field is used as a dimension in the view, it slices the
measure. So instead of an overall sum of sales, the preceding view shows the sum of sales
for each region.

Discrete and continuous

Another important distinction to make with fields is whether a field is being used as
discrete or continuous. Whether a field is discrete or continuous, determines how Tableau
visualizes it based on where it is used in the view. Tableau will give you a visual indication
of the default for a field (the color of the icon in the data pane) and how it is being used in
the view (the color of the active field on a shelf). Discrete fields, such as Region in the
previous example, are blue, and continuous fields, such as Sales, are green.

In the screenshots, in the print version of this book, you should be able to
distinguish a slight difference in shading between discrete (green) and
continuous (blue) fields, but pay special attention to the interface as you
follow along using Tableau.

Discrete fields
Discrete (blue) fields have values that are shown as distinct and separate from each other.
Discrete values can be reordered and still make sense.

[ 13 ]
Other documents randomly have
different content
to a female blossom of the same species. The beginnings of the
modification of the unisexual flowers in this direction may be seen in
variations which occur even now, for we not infrequently find, in a
male catkin, individual blossoms, which, in addition to the stamens,
possess also a pistil with a stigma. (Fig. 50 E shows such an
abnormal hermaphrodite flower from a poplar.)
As soon as hermaphrodite flowers came into existence the struggle
to attract insects began in a more intense degree. Every little
improvement in this direction would form the starting-point of a
process of selection, and would be carried on and increased to the
highest possible pitch of perfection.
It was probably the outer envelopes of the blossoms which first
changed their original green into other colours, usually those which
contrasted strongly with the green, and thus directed the attention
of the insects to the flowers. Variations in the colour of ordinary
leaves are always cropping up from time to time, whether it be that
the green is transformed into yellow or that the chlorophyll
disappears more or less completely and red or blue coloured juices
take its place. Many insects can undoubtedly see colour, and are
attracted by the size of coloured flowers, as Hermann Müller found
by counting the visits of insects to two nearly related species of
mallow, one of which, Malva silvestris, has very large bright rose-red
flowers visible from afar, while the other, Malva rotundifolia, has very
inconspicuous small pale-red flowers. To the former there were
thirty-one different visitors, to the latter he could only make sure of
four. The second species, as is to be expected, depends chiefly on
self-fertilization.
It has recently been disputed from various quarters that insects are
attracted by the colours of the flowers, and these objections are
based chiefly on experiments with artificial flowers. But when, for
instance, Plateau, in the course of such experiments saw bees and
butterflies first fly towards the artificial flowers, and then turn away
and concern themselves no more about them, that only proves that
their sight is sharper than we have given them credit for; for though
they may be deceived at a distance, they are not so when they are
near; it is possible, too, that the sense of smell turns the scale[9]. I
have myself made similar experiments with diurnal butterflies, before
which I placed a single artificial chrysanthemum midst a mass of
natural flowers. It rarely happened indeed that a butterfly settled on
the artificial flower; they usually flew first above it, but did not
alight. Twice, however, I saw them alight on the artificial flower, and
eagerly grope about with the proboscis for a few moments, then fly
quickly away. They had visited the real chrysanthemums or horse-
daisies with evident delight, and eagerly sucked up the honey from
the many individual florets of every flower, and they now
endeavoured to do the same in the artificial flower, and only desisted
when the attempt proved unsuccessful. In this experiment the
colours were of course only white and yellow; with red and blue it is
probably more difficult to give the exact impression of the natural
flower-colours; and in addition there is the absence of the delicate
fragrance exhaled by the flower.
[9] The experiments of Plateau have since been criticized by
Kienitz-Gerloff, who altogether denies their value (1903).

It must be allowed that the colour is certainly not the sole attraction
to the flower; the fragrance helps in most cases, and even this is not
the object of the insect's visits. The real object is the nectar, to
which colour and fragrance only show the way. The development of
fragrance and nectar must, like that of the colour, have been carried
on and increased by processes of selection, which had their basis in
the necessity for securing insect-visits, and as soon as these main
qualities of the flower were established greater refinements would
begin, and flower-forms would be evolved, which would diverge
farther and farther, especially in shape, from the originally simple
and regular form of the blossom.
The reason for this must have lain chiefly in the fact that, after
insect-visits in general were secured by a flower, it would be
advantageous to exclude all insects which would pillage the nectar
without rendering in return the service of cross-fertilization—all
those, therefore, which were unsuited either because of their minute
size or because of the inconstancy of their visits. Before the
butterflies and the bees existed, the regularly formed flat flower with
unconcealed nectar would be visited by a mixed company of caddis-
flies, saw-flies, and ichneumon-flies. But as the nectar changed its
place to the deeper recesses of the flower it was withdrawn from all
but the more intelligent insects, and thus the circle of visitors was
already narrowed to some extent. But when in a particular species
the petals fused into a short tube, all visitors were excluded whose
mouth-parts were too short to reach the nectar; while among those
which could reach it the process of proboscis-formation began; the
under lip, or the first maxillæ, or both parts together, lengthened
step for step with the corolla-tube of the flower, and thus from the
caddis-flies came the butterflies, and from the ichneumon-flies the
burrowing-wasps (Sphegidæ) and the bees.
At first sight one might perhaps imagine that it would have been
more advantageous to the flowers to attract a great many visitors,
but this is obviously not the case. On the contrary, specialized
flowers, accessible only to a few visitors, have a much greater
certainty of being pollinated by them, because insects which only fly
to a few species are more certain to visit these, and above all to visit
many flowers of the same species one after another. Hermann Müller
observed that, in four minutes, one of the humming-bird hawk-
moths (Macroglossa stellatarum) visited 108 different flowers of the
same species, the beautiful Alpine violet (Viola calcarata), one after
the other, and it may have effected an equal number of pollinations
in that short time.
It was, therefore, a real advantage to the flowers to narrow their
circle of visitors more and more by varying so that only the useful
visitors could gain access to their nectar, and that the rest should be
excluded. Thus there arose 'bee-flowers,' 'butterfly-flowers,' 'hawk-
moth flowers,' and, indeed, in many cases, a species of flower has
become so highly specialized that its fertilization can only be brought
about by a single species of insect. This explains the remarkable
adaptations of the orchids and the enormous length of the proboscis
in certain butterflies. Even our own hawk-moths Macroglossa
stellatarum and Sphinx convolvuli show an astonishing length of
proboscis, which measures 8 cm. in the latter species. In Macrosilia
cluentius, in Brazil, the proboscis is 20 cm. in length; and in
Madagascar there grows an orchid with nectaries 30 cm. in length,
filled with nectar to a depth of 2 cm., but the fertilizing hawk-moth is
not yet known.
Thus we may say that the flowers, by varying in one direction or
another, have selected a definite circle of visitors, and, conversely,
that particular insect-groups have selected particular flowers for
themselves, for those transformations of the flowers were always
most advantageous which secured to them the exclusive visits of
their best crossing agents, and these transformations were, on the
one hand, such as kept off unwelcome visitors, and, on the other
hand, such as attracted the most suitable ones.
From the botanical point of view the assumption that flowers and
flower-visiting insects have been adapted to each other by means of
processes of selection has been regarded as untenable, because
every variation in the flower presupposes a corresponding one in the
insect. I should not have mentioned this objection had it not come
from such a famous naturalist as Nägeli, and if it were not both
interesting and useful in our present discussion. Nägeli maintained
that selection could not, for instance, have effected a lengthening of
the corolla-tube of a flower, because the proboscis of the insects
must have lengthened simultaneously with it. If the corolla-tube had
lengthened alone, without the proboscis of the butterfly being at the
same time elongated, the flower would no longer be fertilized at all,
and if the lengthening of the proboscis preceded that of the corolla-
tube it would have no value for the butterfly, and could not therefore
have been the object of a process of selection.
This objection overlooks the facts that a species of plant and of
butterfly consists not of one individual but of thousands or millions,
and that these are not absolutely uniform, but in fact
heterogeneous. It is precisely in this that the struggle for existence
consists—that the individuals of every species differ from one
another, and that some are better, others less well constituted. The
elimination of the latter and the preferring of the former constitutes
the process of selection, which always secures the fitter by
continually rejecting the less fit. In the case we are considering,
then, there would be, among the individuals of the plant-species
concerned, flowers with a longer and flowers with a shorter corolla-
tube, and among the butterflies some with a longer and some with a
shorter proboscis. If among the flowers the longer ones were more
certain to be cross-fertilized than the shorter ones, because hurtful
visitors were better excluded, the longer ones would produce more
and better seeds, and would transmit their character to more
descendants; and if, among the butterflies, those with the longer
proboscis had an advantage, because the nectar in the longer tubes
would, so to speak, be reserved for them, and they would thus be
better nourished than those with the shorter proboscis, the number
of individuals with long proboscis must have increased from
generation to generation. Thus the length of the corolla-tube and
the length of the proboscis would go on increasing as long as there
was any advantage in it for the flower, and both parties must of
necessity have varied pari passu, since every lengthening of the
corolla was accompanied by a preferring of the longest proboscis
variation. The augmentation of the characters depended on, and
could only have depended on, a guiding of the variations in the
direction of utility. But this is exactly what we call, after Darwin and
Wallace, Natural Selection.
We have, however, in the history of flowers, a means of
demonstrating the reality of the processes of selection in two other
ways. In the first place, it is obvious that no other interpretation can
be given of such simultaneous mutual adaptations of two different
kinds of organisms. If we were to postulate, as Nägeli, for instance,
did, an intrinsic Power of Development in organisms, which produces
and guides their variations, we should, as I have already said, be
compelled also to take for granted a kind of pre-established
harmony, such as Leibnitz assumed to account for the correlation of
body and mind: plant and insect must always have been
correspondingly altered so that they bore the same relation to each
other as two clocks which were so exactly fashioned that they
always kept time, though they did not influence each other. But the
case would be more complicated than that of the clocks, because
the changes which must have taken place on both sides were quite
different, and yet at the same time such that they corresponded as
exactly as Will and Action. The whole history of the earth and of the
forms of life must, therefore, have been foreseen down to the
smallest details, and embodied in the postulated Power of
Development.
But such an assumption could hardly lay claim to the rank of a
scientific hypothesis. Although every grain of sand blown about by
the wind on this earth could certainly only have fallen where it
actually did fall, yet it is in the power of any of us to throw a handful
of sand wherever it pleases us, and although even this act of
throwing must have had its sufficient reason in us, yet no one could
maintain that its direction and the places where the grains fell were
predestined in the history of the earth. In other words: That which
we call chance plays a part also in the evolution of organisms, and
the assumption of a Power of Development, predestinating even in
detail, is contradicted by the fact that species are transformed in
accordance with the chance conditions of their life.
This can be clearly demonstrated in the case of flowers. That the
wild pansy (Viola tricolor), which lives in the plains and on
mountains of moderate elevation, is fertilized by bees, and the
nearly allied Viola calcarata of the High Alps by Lepidoptera, is
readily intelligible, since bees are very abundant in the lower region,
and make the fertilization of the species a certainty, while this is not
so in the High Alps. There the Lepidoptera are greatly in the
majority, as every one knows who has traversed the flower-decked
meads of the High Alps in July, and has seen the hundreds and
thousands of butterflies and moths which fly from flower to flower.
Thus the viola of the High Alps has become a 'butterfly-flower' by
the development of its nectaries into a long spur, accessible only to
the proboscis of a moth or butterfly. The chance which led certain
individuals of the ancestral species to climb the Alps must also have
supplied the incentive to the production of the changes adapted to
the visits of the prevalent insect. The hypothesis of a predestinating
Power of Development suffers utter shipwreck in face of facts like
these.
We have, furthermore, an excellent touchstone for the reality of the
processes of selection in the quality of the variations in flowers and
insects. Natural selection can only bring about those changes which
are of use to the possessors themselves; we should therefore expect
to find among flowers only such arrangements as are, directly or
indirectly, of use to them, and, conversely, among insects only such
as are useful to the insect.
And this is what we actually do find. All the arrangements of the
flowers—their colour, their form, their honey-guides, their hairy
honey-paths (Iris), their fragrance, and their honey itself—are all
indirectly useful to the plant itself, because they all co-operate in
compelling the honey-seeking insect to effect the fertilization of the
flower. This is most clearly seen in the case of the so-called
'Deceptive' flowers, which attract insects by their size and beauty,
their fragrance, and their resemblance to other flowers, and force
their visitors to be the means of their cross-fertilization, although
they contain no nectar at all. This is the case, according to Hermann
Müller, with the most beautiful of our indigenous orchids, the lady's
slipper (Cypripedium calceolaris). This flower is visited by bees of the
genus Andrena, which creep into the large wooden-shoe-shaped
under lip in the search for honey, only to find themselves prisoners,
for they cannot get out, at least by the way they came in, because
of the steep and smoothly polished walls of the flower. There is only
one way for the bee; it must force itself under the stigma, which it
can only do with great exertion, and not without being smeared with
pollen, which it carries to the next flower into which it creeps. It can
only leave this one in the same way, and thus the pollen is
transferred to the stigma by a mechanical necessity.
Such remarkable cases remind us in some ways of those cases of
mimicry in which the deceptions have to be used with caution or
they lose their effect. One might be disposed to imagine that such
an intelligent insect as a bee would not be deceived by the lady's
slipper more than once, and would not creep into a second flower
after discovering that there was no nectar in the first. But this
conclusion is not correct, for the bees are well accustomed in many
flowers to find that the nectar has already been taken by other bees;
they could therefore not conclude from one unsuccessful visit that
the Cypripedium did not produce nectar at all, but would try again in
a second, a third, and a fourth flower. If these orchids had
abundantly covered flower-spikes like many species of Orchis, and if
the species were common, the bees would probably soon learn not
to visit them, but the reverse is the case. There is usually only one
or, at most, two open flowers on the lady's slipper, and the plant is
rare, and probably occurs nowhere in large numbers.
If we could find a flower in which the nectar lay open and accessible
to all insects, and which did not require any service from them in
return, the case could not be interpreted in terms of natural
selection; but we do not know of any such case.
Conversely, too, there are no adaptations in the insects which are
useful only to the flowers, and which are not of some use, directly or
indirectly, to the insect itself. Bees and butterflies certainly carry the
pollen from one flower to the stigma of another, but they are not
impelled to do this by a special instinct; they are forced to do it by
the structure of the flower, which has its stamens so placed and
arranged that they must shake their pollen over the visitor, or it may
be that the anthers are modified into stalked, viscid pollinia which
spring off at a touch, and fix themselves, so to speak, on the insect's
head. And even this is not all in the case of the orchis, for the insect
would never of its own accord transfer these pollinia on to the
stigma of the next flower; this is effected by the physical peculiarity
which causes the pollinia, after a short time, to bend forwards on
the insect's head.
All this fits in as well as possible
with the hypothesis: how could an
instinct to carry pollen from one
flower to the stigma of another
have been developed in an insect
through natural selection, since
the insect itself has nothing to
gain from this proceeding?
Accordingly, we never find in the
insect any pincers or any kind of
grasping organ adapted for seizing
and transmitting the pollen.
Fig. 51. The Yucca-moth (Pronuba
There is, however, one very yuccasella). M, laying eggs in
remarkable case in which this the ovary of the Yucca flower.
appears to be so, indeed really is n, the stigma. After Riley.
so, and nevertheless it is not
contradictory to, but is
corroborative of, the theory of selection. The excellent American
entomologist, Riley, established by means of careful observations
that the large white flowers of the Yucca are fertilized by a little
moth which behaves in a manner otherwise unheard of among
insects. Only the females visit the flowers, and they at once busy
themselves collecting a large ball of pollen. To this end they have on
the maxillary palps (Fig. 52, C, mxp) a long process (si), curved in
the form of a sickle, and covered with hairs, which probably no other
Lepidopteron possesses, with the help of which the moth very
quickly sweeps together a ball of pollen, it may be three times the
size of her own head. With this ball the insect flies to the next
flower, and there she lays her egg, by means of an ovipositor
otherwise unknown among Lepidoptera (Fig. 52, A, op), in the pods
of the flower. Finally, she pushes the ball of pollen deep into the
funnel-shaped stigmatic opening on the pistil (Fig. 51, n), and so
effects the cross-fertilization. The ovules develop, and when the
caterpillars emerge from the egg four to five days later they feed on
these until they are ready to enter on the pupa stage. Each little
caterpillar requires about eighteen or twenty seeds for its
nourishment (Fig. 52, B, r).

Fig. 52. The fertilization of the Yucca. A, ovipositor of the

Yucca-moth. op, its sheath. sp, its apex. op1, the
protruded oviduct. B, two ovaries of the Yucca, showing
the holes by which the young moths escape, and (r) a
caterpillar in the interior. C, head of the female moth,
with the sickle-shaped process (si) on the maxillary palps
for sweeping off the pollen and rolling it into a ball. mx1,
the proboscis. au, eye. p1 base of first leg. D, longitudinal
section through an ovary of the Yucca, soon after the
laying of two eggs (ei). stk, the canal made by the
ovipositor.

Here, then, we find an adaptation of certain parts of the moth's body

in relation to the fertilization of the flower, but in this case it is as
much in the interest of the moth as of the plant. By carrying the
pollen to the stigma the moths secure the development of the
ovules, which serve their offspring as food, so that we have here to
do with a peculiar form of care for offspring, which is not more
remarkable than many other kinds of brood-care in insects, such as
ants, bees, Sphex-wasps, ichneumon-flies, and gall-flies.
It might be objected that this case of the Yucca is not so much one
of effecting fertilization as of parasitism; but the eggs, which are laid
in the seed-pods, are very few, and the caterpillars which emerge
from them only devour a very small proportion of the seeds, of
which there may be about 200 (Fig. 52, B). Thus the plants also
derive an advantage from the moth's procedure, for quite enough
seeds are left. The form and position of the stamens and of the
stigma seem to be as exactly adapted to the visits of the moth as
the moth is to the transference of the pollen, for the Yucca can only
be fertilized by this one moth, and sets no seed if the moth be
absent. For this reason the species of Yucca cultivated in Europe
remain sterile.
Thus the apparent contradiction is explained, and the facts
everywhere support the hypothesis that the adaptations between
flowers and insects depend upon processes of selection.
This origin is incontrovertibly proved, it seems to me, in another
way, namely, by the merely relative perfection of the adaptations, or
rather, by their relative imperfection.
I have already pointed out that all adaptations which depend upon
natural selection can only be relatively perfect, as follows from the
nature of their efficient causes, for natural selection only operates as
long as a further increase of the character concerned would be of
advantage to the existence of the species. It cannot be operative
beyond this point, because the existence of the species cannot be
more perfectly secured in this direction, or, to speak more precisely,
because further variations in the direction hitherto followed would no
longer be improvements, even though they might appear so to us.
Thus the corolla of many flowers is suited to the thick, hairy head
and thorax of the bee, for to these only does the pollen adhere in
sufficient quantity to fertilize the next flower; yet the same flowers
are frequently visited by butterflies, and in many of them there has
been no adaptation to prevent these useless visits. Obviously this is
because preventive arrangements could only begin, according to our
theory, when they were necessary to the preservation of the species;
in this case, therefore, only when the pillaging visits of the
butterflies withdrew so many flowers from the influence of the
effective pollinating visitor, the bee, that too few seeds were formed,
and the survival of the species was threatened by the continual
dwindling of the normal number. As long as the bees visit the
flowers frequently enough to ensure the formation of the necessary
number of seeds a process of selection could not set in; but should
the bees find, for instance, that nearly all the flowers had been
robbed of their nectar, and should therefore visit them less diligently,
then every variation of the flower which made honey less accessible
to the butterflies would become the objective of a process of
selection.
Everywhere we find similar imperfections of adaptation which
indicate that they must depend on processes of selection. Thus
numerous flowers are visited by insects other than those which
pollinate them, and these bring them no advantage, but merely rob
them of nectar and pollen; the most beautiful contrivances of many
flowers, such as Glycinia, which are directed towards cross-
fertilization by bees, are rendered of no effect because wood-bees
and humble-bees bite holes into the nectaries from the outside, and
so reach the nectar by the shortest way. I do not know whether
bees in the native land of the Glycinia do the same thing, but in any
case they can do no sensible injury to the species, since otherwise
processes of selection would have set in which would have
prevented the damage in some way or other, whether by the
production of stinging-hairs, or hairs with a burning secretion, or in
some other way. If the actual constitution of the plant made this
impossible, the species would become less abundant and would
gradually die out.
Thus the relative imperfection of the flower-adaptations, which in
general are so worthy of admiration, affords a further indication that
their origin is due to processes of selection.
 ADDITIONAL NOTE TO CHAPTER X.
It has been remarked that the chapter on the Origin of Flowers in
the German Edition contains no discussion and refutation of the
objections which have up till recently been urged against the theory
of flowers propounded by Darwin and Hermann Müller. I admit that
this chapter seemed to be so harmonious and so well rounded, and
at the same time so convincing as to the reality of the processes of
selection, that the feeble objections to it, and the attempts of
opponents to find another explanation of the phenomena, might well
be disregarded in this book.
However, the most important of these objections and counter-
theories may here be briefly mentioned.
Plateau in Ghent was the first to collect facts which appeared to
contradict the Darwinian theory of flowers; he observed that insects
avoided artificial flowers, even when they were indistinguishable in
colour from natural ones as far as our eyes could perceive, and he
concluded from this that it is not the colour which guides the insects
to the flowers, that they find the blossoms less by their sense of
sight than by their sense of smell. But great caution is required in
drawing conclusions from experiments of this kind. I once placed
artificial marguerites (Chrysanthemum leucanthemum) among
natural ones in a roomy frame in the open air, and for a considerable
time I was unable to see any of the numerous butterflies (Vanessa
urticæ), which were flying about the real chrysanthemums, settle on
one of the artificial flowers. The insects often flew quite close to
them without paying them the least attention, and I was inclined to
conclude that they either perceived the difference at sight, or that
they missed the odour of the natural flowers in the artificial ones.
But in the course of a few days it happened twice in my presence
that a butterfly settled on one of the artificial blooms and
persistently groped about with fully outstretched tube to find the
entrance to the honey. It was only after prolonged futile attempts
that it desisted and flew away. That bees are guided by the eye in
their visits to flowers has been shown by A. Forel, who cut off the
whole proboscis, together with the antennæ, from humble-bees
which were swarming eagerly about the flowers. He thus robbed
them of the whole apparatus of smell, and nevertheless they flew
down from a considerable height direct to the same flowers. An
English observer, Mr. G. N. Bulman, has been led to believe, with
Plateau, that it is a matter of entire indifference to the bees whether
the flowers are blue, or red, or simply green in colour, if only they
contain honey, and that therefore the bees could have played no
part in the development of blue flowers, as Hermann Müller
assumed they had, and that they could have no preference for blue
or any other colour, as Sir John Lubbock and others had concluded
from their experiments. This is correct in so far that bees feed as
eagerly on the greenish blossoms of the lime-tree as they do on the
deep-blue gentian of the Alpine meadows or the red blossoms of the
Weigelia, the dog-roses of our gardens or the yellow buttercups
(Ranunculus) of our meadows; they despise nothing that yields them
honey. But it certainly does not follow from this that the bees may
not, under certain circumstances, have exercised a selecting
influence upon the fixation and intensification of a new colour-variety
of a flower. This is less a question of a colour-preference, in the
human sense, on the part of the bees than of the greater visibility of
the colour in question in the environment peculiar to the flower, and
of the amount of rivalry the bees meet with from other insects in
regard to the same flower. In individual cases this would be difficult
to demonstrate, especially since we can form only an approximate
idea of the insect's power of seeing colour, and cannot judge what
the colours of the individual blossoms count for in the mosaic picture
of a flowery meadow. Yet this is the important point, for, as soon as
the bees perceive one colour more readily than another, the
preponderance of this colour-variety over other variations is assured,
since it will be more frequently visited. In the same way we cannot
guess in individual cases why one species of flower should exhale
perfume while a nearly related species does not. But when we
remember that many flowers adapted for the visits of dipterous
insects possess a nauseous carrion-like smell, by means of which
they not only attract flies but scare off other insects, we can readily
imagine cases in which it was of importance to a flower to be able to
be easily found by bees without betraying itself by its pleasant
fragrance to other less desirable visitors.
Thus, therefore, we can understand the odourless but intensely blue
species of gentian, if we may assume that its blue colour is more
visible to bees than to other insects. If I were to elaborate in detail
all the principles which here suggest themselves to me I should
require to write a complete section, and I am unwilling to do this
until I can bring forward a much larger number of new observations
than I am at present in a position to do. All I wish to do here is to
exhort doubters to modesty, and to remind them that these matters
are exceedingly complex, and that we should be glad and grateful
that expert observers like Darwin and Hermann Müller have given us
some insight into the principles interconnecting the facts, instead of
imagining whenever we meet with some little apparently
contradictory fact, which may indeed be quite correct in itself, that
the whole theory of the development of flowers through insects has
been overthrown. Let us rather endeavour to understand such facts,
and to arrange them in their places as stones of the new building.
Often the contradiction is merely the result of the imperfect
theoretical conceptions of its discoverer, as we have already shown
in regard to Nägeli. Bulman, too, fancies he has proved that bees do
not distinguish between the different varieties of a flower, but visit
them indiscriminately with the same eagerness, thus causing
intercrossing of all the varieties, and preventing any one from
becoming dominant. But are the varieties which we plant side by
side in our gardens of the kind that are evolved by bees? That is to
say, are their differences such as will turn the scale for or against the
visits of the bees? If one were less, another more easily seen by the
bees; or if one were more fragrant, or had a fragrance more
agreeable to bees than the other, the result of the experiment would
probably have been very different.
One more objection has been made. It is said that the bees,
although exclusively restricted, both themselves and their
descendants, to a diet of flowers, are not so constant to a particular
flower as the theory requires. They do indeed exhibit a 'considerable
amount of constancy,' and often visit a large number of flowers of
the same species in succession, but the theory requires that they
should not only confine themselves to this one species, but to a
single variety of this species. These views show that their authors
have not penetrated far towards an understanding of the nature of
selection. Nature does not operate with individual flowers, but with
millions and myriads of them, and not with the flowers of a single
spring, but with those of hundreds and thousands of years. How
often a particular bee may carry pollen uselessly to a strange flower
without thereby lowering the aggregate of seeds so far that the
existence of the species seems imperilled, or how often she may
fertilize the pistil of a useful variation with the pollen of the parent
species, without interrupting or hindering the process of the
evolution of the variety, no mortal can calculate, and what the
theory requires can only be formulated in this way: The constancy of
the bees in their visits to the flowers must be so great that, on an
average, the quantity of seeds will be formed which suffices for the
preservation of the species. And in regard to the transformation of a
species, the attraction which the useful variety has for the bees
must, on an average, be somewhat stronger than that of the parent
species. As soon as this is the case the seeds of the variety will be
formed in preponderant numbers, although they may not all be quite
pure from the first, and by degrees, in the course of generations, the
plants of the new variety will preponderate more and more over
those of the parent form, and finally will alone remain. In the first
case we have before our eyes the proof that, in spite of the
imperfect constancy of the bees, a sufficient number of seeds is
produced to secure the existence of the species. Or does Mr. Bulman
conclude from the fact that the bees are not absolutely constant that
flowers are not fertilized by bees at all?
I cannot conclude this note without touching briefly upon what the
opponents of the flower theory have contributed, and what
explanation of the facts they are prepared to offer.
In his important work, Mechanische-physiologische Theorie der
Abstammungslehre, published in 1884, Nägeli, as a convinced
opponent of the theory of selection, attempted an explanation. He
was quite aware that his assumption of an inward 'perfecting
principle' would not suffice to explain the mutual adaptations of
flowers and insects, and he refers the transformation of the first
inconspicuous blossoms into flowers to the mechanical stimulus
which the visiting insects exerted upon the parts of the blossom. By
the pressure of their footsteps, the pushing and probing with their
proboscis, they have, he says, transformed gradually, for instance,
the little covering leaves at the base of a pollen vessel into large
flower petals, caused the conversion of short flower-tubes into long
ones, and of the pollen, once dry and dusty, into the firmly adhesive
mass formed in the anther lobes of our modern flowers. The colour
of the flowers depends, according to him, upon the influence of
light, which certainly no more explains the yellow ring on a blue
ground in the forget-me-not than it does the many other nectar-
guides which show the insect the way to the honey. Nägeli works
with the Lamarckian principle in the most daring way, and with the
same naïveté as Lamarck himself in his time, that is, without offering
any sort of explanation as to how the minute impression made, say
by the foot or by the proboscis of an insect, upon a flower, is to be
handed on to the flowers of succeeding generations. He treats the
unending chain of generations as if it were a single individual, and
operates with his 'secular' stimulus, and with 'weak stimuli, lasting
through countless generations,' as though they were a proved fact.
But I have not even touched upon the question as to whether these
'stimuli' could produce the changes he ascribes to them, even if they
were continually affecting the flower. How the scale-like covering
leaves of the pollen vessels could become larger and petal-like
through the treading of an insect's foot is as difficult to see as why a
honey-tube should become longer because of the butterfly's honey-
sucking: might it not just as well become wider, narrower, or even
shorter? I see no convincing reason why it should become longer!
And even if it did so, it would necessarily continue to lengthen as
time went on, and this is not the case, for we find corolla-tubes of all
possible lengths, but, it is to be noted, always in harmony with the
length of the proboscis of the visiting insect. In a similar way
Henslow has recently attempted to refer the origin of flowers to the
mechanical stimulus exercised upon it by the visiting insects. 'An
insect hanging to the lower petal of a flower elongates the same by
its weight, and the lengthened petal is transmitted by
heredity.'...'The irritation caused by its feet in walking along the
flower causes the appearance of colouring matter, and the colour is
likewise transmitted.'...'As it probes for honey it causes a flow of
sweet sap to that part, and this also becomes hereditary!'
In this case, also, it is simply taken for granted that every little
passing irritation not only produces a perceptible effect, but that this
effect is transmissible. In a later lecture we shall have to discuss in
detail the question of the inheritance of functional modifications. It is
enough to say here that, if this kind of transmission really took place
even in the case of such minute and transitory changes, there could
be no dispute as to the correctness of the 'Lamarckian principle,'
since every fairly strong and lasting irritation could be demonstrated
with certainty to produce an effect. When a butterfly, floating freely
in the air, sucks honey from a tube, the irritation must be almost
analogous to that caused by a comb lightly drawn by some one
through our hair, and this is supposed to effect the gradual
lengthening of the corolla-tube of the flower!
The secretion of honey, too, depends upon the persistent irritation of
the proboscis! Then 'deceptive flowers,' like the Cypripedium we
have mentioned, could not exist at all, for they contain no honey,
although the proboscis of the bee must cause the same irritation in
them as in other orchids which do contain honey. This whole 'theory'
of direct effect is, moreover, only a crude and apparent
interpretation, which explains the conditions only in so far as they
can be seen from a distance; it fails as soon as they are more
exactly examined; all the great differences in the position of the
honey, its concealment from intelligent insects, its protection from
rain by means of hairs, and against unwelcome guests by a sticky
secretion, the development of a corolla-tube which corresponds in
length to the length of the visiting insect's proboscis, the
development of spurs on the flower, in short, all the numerous
contrivances which have reference to cross-fertilization by insects
remain quite unintelligible in the light of this theory—it is a mere pis
aller explanation for those who continue to struggle against
accepting the theory of selection.
 LECTURE XI
SEXUAL SELECTION
Decorative colouring of male butterflies and birds—Wallace's
interpretation—Preponderance of males—Choice of the females—Sense
by sight in butterflies—Attractive odours—Scent-scales—Fragrance of
the females—The limits of natural and sexual selection not clearly
defined—Odours of particular species—Odours of other animals at the
breeding season—Song of the Cicadas, and of birds—Diversity of
decoration successively acquired—Humming-birds—Substitution of
other aids to wooing in place of personal decoration—Smelling organs
of male insects and crabs—Contrivances for seizing and holding the
female—Small size of certain males—Weapons of males used in
struggle for the females—Turban eyes of Ephemerids—Hoods that can
be inflated on the head of birds—Absence of secondary sexual
characters in lower animals—Transference of male characters to the
females—Lycæna—Parrots—Fashion operative in the phyletic
modifications of colour—Pattern of markings on the upper surface of a
butterfly's wing simpler than on the under side—Conclusion.

We found in the process of Natural Selection an explanation of

numerous effective adaptations in plants and animals, as regards
form, colouring, and metabolism, of the most diverse weapons and
protective devices, of the existence of those forms of blossoms
which we call flowers, of instincts, and so on. The origin of the most
characteristic parts of whole orders of insects can only be
understood as adaptations to the environment brought about by
means of natural selection. Impressed by this, we have now to ask
whether all the transformations of organisms may not be referred to
adaptation to the continually changing conditions of life? We shall
return to this question later, but in the meantime we are far from
being able to answer it in the affirmative, for there are undoubtedly
a great many characters, at least in animals, which cannot have
owed their origin to natural selection in the form in which we have
studied it so far.
How could the splendid plumage of the humming-birds, of the
pheasants, of the parrots, the wonderful colour-patterns of so many
diurnal butterflies, be referred to the process of natural selection,
since all these characters can have no significance for their
possessors in the struggle for existence? Or of what use in the
struggle for existence could the possession of its gorgeous dress of
feathers be to the bird of Paradise; or of what service is the azure
blue iridescence of the Morpho of Brazil, which makes it conspicuous
from a distance when it plays about the crowns of the palm-trees?
We might indeed suppose that they are warning signs of
unpalatableness, like those of the Heliconiides or of the gaily
coloured caterpillars, but, in the first place, these gay creatures are
by no means inedible, and are indeed much persecuted; and,
secondly, the females have quite different and very much darker and
simpler colours. The gleaming splendour of all these birds of
Paradise and humming-birds, as well as that of many butterflies, is
found in the male sex only. The females of the birds just mentioned
are dark in colour and without the sparkling decorative feathers of
the males; they are plain—just like the females of many butterflies.
Alfred Russel Wallace has suggested that the explanation of this lies
in the greater need of the females for protection, since, as is well
known, they usually perform the labours of brooding, and are thus
frequently exposed to the attacks of enemies. It is undoubtedly true
that the dark and inconspicuous colouring of many birds and
butterflies depends on this need for protection, but this does not
explain the brilliant colours of the males of these species. Or can it
be that these require no explanation further than that they are, so to
speak, a chance secondary outcome of the structural relations of the
feathers and wing-scales respectively, which brought with it some
other advantage not known to us? Perhaps something in the same
way as the red colour of the blood in all vertebrates, from fishes
upwards, cannot be useful on the ground that it appears red to us,
but because it is the expression of the chemical constitution of the
hæmoglobin, a body which is indispensable to the metabolism,
which here has the secondary and intrinsically quite unimportant
peculiarity of reflecting the red rays of light.
No one can seriously believe this in regard to butterflies who knows
that their colours are dependent on the scales which thickly cover
the wings, and the significance of which, in part at least, is just to
give this or that colour to the wing. They are degenerate or
colourless among the transparent-winged butterflies, and their
colour depends partly on pigment, partly on fluorescence and
interference conditioned by the fine microscopical structure of a
system of intercrossing lines on faintly coloured scales. The scales of
our male 'blue' butterflies (Lycæna) only appear blue because of
their structure, while the brown scales of their mates are due to a
brown pigment. If the pigment be removed from the scales of the
female by boiling with caustic potash, and they be then dried, they
do not look blue like those of the male; the scales of the male,
therefore, must possess something which those of the female do
not.
Still less will any one be disposed to regard the marvellous splendour
of the plumage of the male bird of Paradise, with its erectile collars—
glistening like burnished metal—on the neck, breast or shoulders,
with its tufts, with its specially decorative feathers standing singly
out from the rest of the plumage, on head, wings, or tail, with its
mane-like bunch of loose, pendulous feathers on the belly and on
the sides, in short, with its extraordinary, diverse, and unique
equipment of feathers, as a mere unintentional accessory effect of a
feather dress designed for flight and protective warmth. Such
conspicuous, diverse, and unusual specializations of plumage must
have some other significance than that just indicated.
Alfred Russel Wallace regards these distinctive features of the male
as an expression of the greater vigour, and the more active
metabolism of the males, but it is unproved that the vigour of the
male birds is greater than that of the females, and it is not easy to
see why a more active metabolism should be necessary for the
production of strikingly bright colours than for that of a dark or
protective colour. Moreover, there are brilliantly coloured females,
both among birds and butterflies, and in nearly allied species the
males may be either gorgeous or quite plain like the females.
Darwin refers the origin of these secondary sexual characters to
processes of selection quite analogous to those of ordinary natural
selection, only that in this case it is not the maintenance of the
species which is aimed at, but the attainment of reproduction by the
single individual. The males are to some extent obliged to struggle
for the possession of the females, and every little variation which
enables a male to gain possession of a female more readily than his
neighbour has for this reason a greater likelihood of being
transmitted to descendants. Thus, attractive variations which once
crop up will be transmitted to more and more numerous males of
the species, and among these it will always be those possessing the
character in question in the highest degree which will have the best
chance of securing a mate, and so the character will continue to be
augmented as long as variations in this direction appear.
Two kinds of preliminary conditions, however, must be assumed. As
the ordinary natural selection could never have operated but for the
fact that in every generation a great many individuals, indeed the
majority of them, perish before they have had time to reproduce, so
the process of sexual selection could never have come into operation
if every male were able ultimately to secure a mate, no matter what
degree of attractiveness to the latter he possessed. If the numbers
of males and females were equal, so that there was always one
female to one male, there could be no choice exercised either by
male or female, for there would always remain individuals enough of
both sexes, so that no male need remain unmated.
But this is not the case: the proportions of the sexes are very rarely
as 1 : 1; there is usually a preponderating number of males, more
rarely of females. Among birds the males are usually in the majority,
still more so among fishes; and among diurnal butterflies there are
often a hundred males to one female (Bates), although there seem
to be a few tropical Papilionidæ among which the females have
rather the preponderance. Darwin called attention to the fact that
one could infer the greater rarity of the females even from the
pricelists of butterflies issued by the late Dr. Staudinger in connexion
with his business, for the females in most species, except the very
common ones, are priced much higher than the males, often twice
as high. In the whole list of many thousands of species there are
only eleven species of nocturnal Lepidoptera in which the males are
dearer than the females.
Among the Mayflies or Ephemerides, too, the males are in the
majority; in many of them there are sixty males to one female: but
there are other kinds of insects, such as the dragon-flies
(Libellulidæ), in which the females are three or four times as
numerous. There are also, it may be remembered, some kinds of
insects, such as Aphides, which have become capable of
parthenogenetic reproduction, and in which the males are becoming
extinct, e.g. in the case of Cerataphis in British orchid-houses.
The first postulate implied in 'sexual selection,' namely, that there be
an unequal number of individuals in the two sexes, is therefore
fulfilled in Nature; we have now to inquire whether the second
condition postulated—the power of choice—may also be regarded as
a reality.
This point has been disputed from many sides, and even by one of
the founders of the whole selection theory, Alfred Russel Wallace.
This naturalist doubts whether a choice is exercised among birds by
either sex in regard to pairing, and maintains that, even if there
could be a choice, this could not have produced such differences in
colour and character of the plumage, since that would presuppose
the existence of similar taste in the females through many
generations. In a similar way it has been doubted whether butterflies
can be said to exercise any real power of sexual choice, whether a
more beautiful male is as such preferred to a less beautiful suitor.
It must be admitted that direct observation of choosing is difficult,
and that as yet there is very little that can be said with certainty on
this point. But there are, after all, some precise observations on
mammals and birds which prove that the female shows active
inclination to, or disinclination for, a particular male. If we hold fast
to this fact, and add to it that the distinctive markings of the males
are wonderfully developed during the period of courtship, and are
displayed before the females, and that they only appear in
mammals, birds, amphibians, and fishes at the time of sexual
maturity, it seems to me that there can be no doubt that they are
intended to fascinate the females, and to induce them to yield
themselves to the males. The opponents of the theory of sexual
selection attach too much importance to isolated cases; they
imagine that each female must make a choice between several
males. But the theory of sexual selection does not demand this, any
more than the theory of natural selection requires the assumption
that every individual of a species which is better equipped for the
struggle for existence must necessarily survive and attain to
reproduction, or, conversely, that the less well equipped must
necessarily perish.
All that the theory requires is, that the selective and eliminative
processes do, on an average, secure their ends, and in the same
way the theory of sexual selection does not need the assumption
that every female is in a position to exercise a scrupulous choice
from among a troop of males, but only that, on an average, the
males more agreeable to the females are selected, and those less
agreeable rejected. If this is the case, it must result in the male
characters most attractive to the females gaining preponderance,
and becoming more and more firmly established in the species,
increasing in intensity, and finally becoming a stable possession of all
the males.
When we go more into details we shall see that the particular
qualities of the distinctive masculine characters are exactly such as
they would be if they owed their existence to processes of selection;
in other words, from this point of view the phenomena of the
decorative sexual characters can be understood up to a certain
point. It seems to me that we are bound to accept the process of
sexual selection as really operative, and instead of throwing doubt
upon it, because the choice of the females can rarely be directly
established, we should rather deduce from the numerous sexual
characters of the males, which have a significance only in relation to
courtship, that the females of the species are sensitive to these
distinguishing characters, and are really capable of exercising a
choice.
In my mind at least there remains no doubt that the 'sexual
selection' of Darwin is an important factor in the transformation of
species, even if I only take into consideration those secondary sexual
characters which are related to wooing. We shall see, however, that
there are others in regard to whose origin through processes of
selection doubt is still less legitimate, and from which, on this
account, we can argue back to the courtship characters.
The first beginning of transformation is not, even in ordinary natural
selection, to be understood as due to selection, but is to be regarded
as a given variation (the causes of which we shall discuss later on);
it is only the increase of such incipient variations in a definite
direction that can depend on natural selection, and they must
depend on it in so far as the transformations are purposeful. Now, all
secondary sexual characters can be recognized as useful, save only
the decorative distinctions, although these also undoubtedly
represent intensifications of originally unimportant variations. Are we
then to regard these alone as the mere outcome of the internal
impulsive forces of the organism, while in the case of the analogous
sexual characters for tracking, catching, and holding the female, and
so forth, the augmentation and the directing must be referred to
processes of selection? But if there be any utility at all in the
decorative sexual characters it can only lie in their greater
attractiveness to the females, and it can only be of any account if
the females have, in a certain sense, the power of choice.
Independently, therefore, of direct observations as to the actual
occurrence of choosing, we should be compelled by our chain of
reasoning to assume that there was such a power of choice—and I
shall immediately discuss it more precisely.
If we consider the decorative, distinctive characters of the males
more closely, we find that they are of very diverse kinds. The males
of many animals are distinguished from the females chiefly by
greater beauty of form, and especially of colour. This is the case in
many birds, some amphibians, like the water-salamander, many
fishes, many insects, and above all, in diurnal Lepidoptera. Especially
among birds the dimorphism between the sexes is in obvious
relation to the excess in the number of male individuals, or—what
practically comes to the same thing—to polygamy. For when a male
attaches to himself four or ten females the result is the same as if
the number of female individuals were divided by four or by ten.
Thus the fowls and pheasants, which are polygamous, are adorned
by magnificent colours in the male sex, while the monogamous
partridges and quails exhibit the same colouring in both sexes. Of
course 'beautiful' is a relative term, and we must not simply assume
that what seems beautiful to us appears so to all animals; yet when
we see that all the male birds which are beautifully decorative
according to our taste—whether humming-birds, pheasants, birds of
Paradise, or rock-cocks (Rupicola crocea)—unfold their 'feather-
wheels, 'fans,' 'collars,' and so forth, before the eyes of the females
in the breeding season, and display them in all their brilliance, we
must conclude that, in these instances at least, human taste accords
with that of the animals. That birds have sharp vision and distinguish
colours is well known; it is not for nothing that the service berries
and many other berries suitable for birds are red, the mistletoe
berries white, in contrast to the evergreen foliage of this plant, the
juniper berries black so that they stand out amid the snows of
winter; in this direction, then, there is no difficulty in the way of
sexual selection.
Even among much lower animals, like the butterflies, there seems to
me no reason for the assumption that they do not see the gorgeous
colours and often very complicated markings, the bars and eye-
spots, on the wings of their fellows of the same species. Of course if
each facet of the insect eye contributed only a single visual
impression, as Johannes Müller supposed, then even an eye with
12,000 facets would give but a rough and ill-defined picture of
objects more than a few feet away, and I confess that for a long
time I regarded this as an obstacle in the way of referring the sexual
dimorphism of butterflies to processes of selection. But we now
know, through Exner, that this is not the case; we know that each
facet gives a little picture, and not an 'inverted' but an 'upright' one,
and experiment with the excised insect eye has directly shown that it
throws on a photographic plate a tolerably clear image of even
distant objects, such as the frame of a window, a large letter painted
on the window, or even a church tower visible through it.
Furthermore, the structure of the eye allows of incomparably clearer
vision of near objects, for in that case the eyes act like lenses, and
reveal much more minute details than we ourselves are able to make
out. Here again, therefore, there is no obstacle to the Darwinian
hypothesis of a choice on the part of the females, for although it
cannot be demonstrated from the structure of the eye itself that
insects see colour, and that colours have a specially exciting
influence on them, yet we can deduce this with certainty from the
phenomena of their life. The butterflies fly to gaily coloured flowers,
and as they find in them their food, the nectar of the flowers, we
may take for granted that the sight of the colour of their food-
providing plants is associated with an agreeable sensation, and this
is an indication that similar colours in their fellows may awaken
similar agreeable sensations.
 Fig. 53. Scent-scales of diurnal butterflies. a, of Pieris. b,
of Argynnis paphia. c, of a Satyrid. d, of Lycæna. All highly
magnified.

This conclusion is furthermore confirmed by the fact that, in the

male sex, numerous species of butterfly possess another means of
exciting the females, namely, by pleasant odours. Volatile ethereal
oils are secreted by certain cells of the skin, and exhale into the air
through specially constructed scales. Usually the apparatus for
dispersing fragrance occurs on the wing in the form of the so-called
scent-scales (Duftschuppen), peculiar modifications of the ordinary
colour-scales of the wing, but sometimes they take the form of
brush-like hair-tufts on the abdomen, and they are in all cases so
arranged that the volatile perfume from the cells of the skin
penetrates into them, and then evaporates through very thin spots
on the surface of the scale, or through brush-like, expanded fringes
on their tips. Many of these have long been known to entomologists,
because their divergence in form from the ordinary scales attracted
attention; and it was also observed that they never occurred on the
females, but only on the males. Their significance, however,
remained obscure until, by a happy chance, Fritz Müller, in his
Brazilian garden, discovered the fact that there are butterflies which
give off fragrance like a flower, and then close investigation revealed
to him the connexion between this delicate odour and the so-called
'male scales.' One can convince oneself of the correctness of the
observation even in some of our own butterflies by brushing the
finger over the wing of a newly caught male Garden White (Pieris
napi). The finger will be found covered with a white dust, the
rubbed-off wing-scales, and it will have a delicate perfume of lemon
or balsam, thus proving that the fragrance adheres to the scales.
In the last case, that is,
among the Whites
(Pieridæ) (Fig. 53, a), the
scent-scales are
distributed fairly regularly
Fig. 54. A portion of the upper surface
over the upper surface of
of the wing of a male 'blue' (Lycæna the wing, and the same is
menalcas); after Dr. F. Köhler. bl, ordinary true of our blue
blue scales. d, scent-scales. Highly butterflies, the Lycænnidæ
magnified. whose minute lute-shaped
scales are shown singly in
Fig. 53, d, but in their natural position among the ordinary scales in
Fig. 54. In many other diurnal, and also in nocturnal Lepidoptera,
the fragrant scales are united into tufts and localized in definite
areas. They then often form fairly large spots, stripes, or brushes,
which are easily visible to the naked eye. Thus the males of our
various species of grass-butterflies (Satyridæ) have velvet-like black
spots on the anterior wings, while the fritillary, Argynnis paphia, has
coal-black stripes on four longitudinal ribs of the anterior wing which
are absent in the females, and which are composed of hundreds of
odoriferous scales. Certain large forest butterflies of South America,
resembling our Apatura, bear in the middle of the gorgeous green
shimmering posterior wing a thick expansible brush of long, bright
yellow scent-scales, and a similar arrangement obtains in the
beautiful violet butterfly of the Malay Islands, the Zeuxidia wallacei
depicted in Fig. 55. In many of the Danaides, which we have already
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