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Contents
1. Preface xi

1. Part I Introduction 1

1. 1 Introduction to Scientific Research 1

1. Traditional Methods of Acquiring Knowledge 2

1. Authority 3

2. Rationalism 3

3. Empiricism 3

2. The Scientific Approach to Knowledge Generation 4

1. Induction and Deduction 4

2. Hypothesis Testing 5

3. Science in the 21st Century 6

3. Assumptions Underlying Scientific Research 7

1. Regularity in Nature 7

2. Reality in Nature 7

3. Discoverability 8

4. Characteristics of Scientific Research 8

1. Empirical 8

2. Control 9
3. Operationalization of Constructs 9

4. Replication 10

5. Evidence Not Proof 10

5. The Role of Theory in Scientific Research 11

6. Objectives of Psychological Research 13

1. Description 13

2. Explanation 13

3. Prediction 13

4. Control or Influence 14

7. Pseudoscience 14

1. Summary 15

2. Key Terms and Concepts 16

3. Related Internet Sites 16

4. Practice Test 16

5. Challenge Exercises 17

2. 2 Research Approaches and Methods of Data Collection 18

1. Variables in Quantitative Research 19

2. Experimental Research 21

3. Causation 22

1. Cause 22
2. Effect 23

3. Required Criteria for Making the Claim of Causation


23

4. The Psychological Experiment 24

1. Example of an Experiment and Its Logic 24

2. Experimental Research Settings 26

3. Advantages of Experimental Research 26

1. 1. Causal Inference 26

2. 2. Ability to Manipulate Variables 27

3. 3. Control over Extraneous Variables 27

4. Disadvantages of Experimental Research 27

1. 1. Does Not Test Effects of Nonmanipulated


Variables 27

2. 2. Artificiality 27

3. 3. Incomplete Method of Scientific Inquiry 27

5. Nonexperimental Quantitative Research 28

1. Advantages and Disadvantages of Nonexperimental


Quantitative Research 30

6. Qualitative Research 31

7. Mixed Methods Research 33

8. Major Methods of Data Collection 33


1. Tests 34

2. Questionnaires 34

3. Interviews 35

4. Focus Groups 35

5. Observation 36

6. Existing or Secondary Data 37

1. Summary 39

2. Key Terms and Concepts 40

3. Related Internet Sites 40

4. Practice Test 40

5. Challenge Exercises 41

2. Part II Planning the Research Study 42

1. 3 Identifying Research Questions and Forming Hypotheses 42

1. Sources of Research Ideas 43

1. Everyday Life 43

2. Practical Issues 43

3. Past Research 44

4. Theory 44

2. Breadth of Research Ideas 45

1. Ideas Not Capable of Scientific Investigation 46


3. Review of the Literature 46

1. Getting Started 47

2. Defining Objectives 47

3. Conducting the Search 47

1. Books 48

2. Psychological Journals 48

3. Computerized or Electronic Databases 48

4. Internet Resources 51

4. Obtaining Resources 53

5. Additional Information Sources 53

4. Feasibility of the Study 54

5. Purpose of the Research 55

1. Defining the Research Question 55

2. Specificity of the Research Question 55

6. Formulating Hypotheses 56

1. Summary 57

2. Key Terms and Concepts 58

3. Related Internet Sites 58

4. Practice Test 58

5. Challenge Exercises 59
2. 4 Ethics 60

1. Research Ethics: What Are They? 61

1. Relationship Between Society and Science 61

2. Professional Issues 62

3. Treatment of Research Participants 64

2. Ethical Dilemmas 64

3. Ethical Guidelines 66

1. Beneficence and Nonmaleficence 67

2. Fidelity and Responsibility 67

3. Integrity 68

4. Justice 68

5. Respect for People’s Rights and Dignity 68

4. APA Ethical Standards for Research 68

1. Institutional Approval 69

2. Informed Consent 70

1. Informed Consent and Minors 70

2. Passive Versus Active Consent 71

3. Deception 72

4. Debriefing 73

5. Coercion and Freedom to Decline Participation 74


6. Confidentiality, Anonymity, and the Concept
of Privacy 75

5. Ethical Issues in Electronic Research 76

1. Informed Consent and Internet Research 76

2. Privacy and Internet Research 77

3. Debriefing and Internet Research 77

6. Ethical Issues in Preparing the Research Report 78

1. Authorship 78

2. Writing the Research Report 79

7. Ethics of Animal (Nonhuman) Research 80

1. Safeguards in the Use of Animals 80

2. Animal Research Guidelines 80

1. I. Justification of the Research 80

2. II. Personnel 81

3. III. Care and Housing of Animals 81

4. IV. Acquisition of Animals 81

5. V. Experimental Procedures 81

6. VI. Field Research 82

7. VII. Educational Use of Animals 82

1. Summary 83

2. Key Terms and Concepts 84


3. Related Internet Sites 85

4. Practice Test 85

5. Challenge Exercises 85

3. Part III Foundations of Research 87

1. 5 Measuring Variables and Sampling 87

1. Defining Measurement 88

1. Scales of Measurement 88

1. Nominal Scale 88

2. Ordinal Scale 89

3. Interval Scale 89

4. Ratio Scale 89

2. Psychometric Properties of Good Measurement 90

1. Overview of Reliability and Validity 90

2. Reliability 90

1. Test–Retest Reliability 90

2. Equivalent-Forms Reliability 90

3. Internal Consistency Reliability 91

4. Interrater Reliability 91

3. Validity 91

1. Validity Evidence Based on Content 92


2. Validity Evidence Based on Internal Structure 93

3. Validity Evidence Based on Relations to Other


Variables 93

4. Using Reliability and Validity Information 95

5. Sources of Information About Tests 95

3. Sampling Methods 95

1. Terminology Used in Sampling 96

2. Random Sampling Techniques 97

1. Simple Random Sampling 97

2. Stratified Random Sampling 99

3. Cluster Random Sampling 101

4. Systematic Sampling 101

3. Nonrandom Sampling Techniques 102

4. Random Selection and Random Assignment 103

5. Determining the Sample Size When Random Sampling Is


Used 104

6. Sampling in Qualitative Research 105

1. Summary 106

2. Key Terms and Concepts 107

3. Related Internet Sites 107

4. Practice Test 108


5. Challenge Exercises 108

2. 6 Research Validity 109

1. Overview of Four Major Types of Validity 110

2. Statistical Conclusion Validity 110

3. Construct Validity 111

1. Threats to Construct Validity 112

1. Participant Reactivity to the Research Situation


112

2. Researcher Effects 113

4. Internal Validity 115

1. Threats to Internal Validity 116

1. History 118

2. Maturation 120

3. Instrumentation 120

4. Testing 120

5. Regression Artifact 121

6. Attrition 121

7. Selection 122

8. Additive and Interactive Effects 122

5. External Validity 123

1. Population Validity 124


2. Ecological Validity 125

3. Temporal Validity 125

4. Treatment Variation Validity 125

5. Outcome Validity 126

6. Relationship Between Internal and External Validity 126

1. Summary 127

2. Key Terms and Concepts 128

3. Related Internet Sites 128

4. Practice Test 128

5. Challenge Exercises 129

3. 7 Control Techniques in Research 131

1. Control Techniques Carried Out at the Beginning of the


Study 133

2. Random Assignment 133

3. Matching 136

1. Matching by Holding Variables Constant 136

2. Matching by Equating Participants 137

4. Building the Extraneous Variable into the Design 138

5. Statistical Control 139

6. Control Techniques Carried Out During the Study 139

7. Counterbalancing 139
1. Randomized Counterbalancing 141

2. Complete Counterbalancing 141

3. Incomplete Counterbalancing 142

8. Blind Technique 143

1. Double-Blind Technique 144

2. Single-Blind Technique 144

3. Partial-Blind Technique 144

9. Automation 145

10. Likelihood of Achieving Control 145

1. Detecting Participant Interpretation 145

1. Summary 146

2. Key Terms and Concepts 147

3. Related Internet Sites 147

4. Practice Test 147

5. Challenge Exercises 148

4. 8 Procedure for Conducting a Research Study 149

1. Institutional Approval 150

2. Research Participants 151

1. Obtaining Animals 151

2. Obtaining Human Participants 151


3. Sample Size 153

1. Power 153

4. Apparatus and/or Instruments 156

5. Procedure 157

1. Scheduling of Research Participants 157

2. Consent to Participate 158

3. Instructions 159

4. Data Collection 159

5. Debriefing 159

1. Debriefing Functions 159

2. How to Debrief 160

6. Pilot Study 161

1. Summary 162

2. Key Terms and Concepts 163

3. Related Internet Site 163

4. Practice Test 163

5. Challenge Exercise 164

4. Part IV Experimental Methods 165

1. 9 Experimental Research Design 165

1. Weak Experimental Research Designs 167


1. One-Group Posttest-Only Design 168

2. One-Group Pretest–Posttest Design 168

3. Posttest-Only Design with Nonequivalent Groups 169

2. Strong Experimental Research Designs 170

3. Between-Participants Designs 172

1. Posttest-Only Control-Group Design 172

1. Strengths and Weaknesses of the Posttest-Only


Control-Group Design 173

4. Within-Participants Designs 175

1. Strengths and Weaknesses of Within-Participants


Designs 175

5. Mixed Designs (i.e., Combination of Between and Within)


176

1. Pretest–Posttest Control-Group Design 177

2. Advantages and Disadvantages of Including a Pretest


177

6. Factorial Designs 179

1. Factorial Designs Based on Within-Participants


Independent Variables 184

2. Factorial Designs Based on a Mixed Model 185

3. Nature of Independent Variables in Factorial Designs


186

4. Strengths and Weaknesses of Factorial Designs 186


7. How to Choose or Construct the Appropriate Experimental
Design 188

1. Summary 189

2. Key Terms and Concepts 190

3. Related Internet Sites 190

4. Practice Test 190

5. Challenge Exercises 191

2. 10 Quasi-Experimental Designs 193

1. Quasi-Experimental Designs 194

2. Nonequivalent Comparison Group Design 196

1. Outcomes with Rival Hypotheses 198

1. Outcome I: Increasing Control and Experimental


Groups 198

2. Outcome II: Experimental-Group-Higher-than-


Control-Group-at-Pretest Effect 199

3. Outcome III: Experimental-Group-Lower-than-


Control-Group-at-Pretest Effect 199

4. Outcome IV: Crossover Effect 200

2. Ruling Out Threats to the Nonequivalent Comparison


Group Design 200

3. Causal Inference from the Nonequivalent Comparison


Group Design 202

3. Time-Series Design 202


1. Interrupted Time-Series Design 203

4. Regression Discontinuity Design 204

1. Summary 206

2. Key Terms and Concepts 207

3. Related Internet Sites 207

4. Practice Test 207

5. Challenge Exercises 208

3. 11 Single-Case Research Designs 210

1. History of Single-Case Designs 211

2. Single-Case Designs 213

1. ABA and ABAB Designs 214

2. Combination Design 216

3. Multiple-Baseline Design 217

4. Changing-Criterion Design 218

3. Methodological Considerations in Using Single-Case


Designs 220

1. Baseline 220

2. Changing One Variable at a Time 221

3. Length of Phases 222

4. Criteria for Evaluating Change 222

1. Experimental Criterion 223


2. Therapeutic Criterion 224

5. Rival Hypotheses 224

1. Summary 225

2. Key Terms and Concepts 226

3. Related Internet Sites 226

4. Practice Test 226

5. Challenge Exercises 227

5. Part V Nonexperimental, Qualitative, and Mixed Methods Research


228

1. 12 Nonexperimental Quantitative Research Methods 228

1. Independent Variables and Dependent Variables


in Nonexperimental Quantitative Research 229

2. The Bradford Hill Criteria for Causation 230

3. Techniques of Control in Nonexperimental Quantitative


Research 232

1. Matching 232

2. Holding the Extraneous Variable Constant 234

3. Statistical Control 234

4. Nonexperimental Designs Based on the Time Dimension


235

1. Cross-Sectional and Longitudinal Designs 235


2. Evaluation of Cross-Sectional and Longitudinal
Designs 237

5. Nonexperimental Research Designs based on Research


Purpose 238

1. Descriptive Nonexperimental Quantitative Research


238

2. Predictive Nonexperimental Quantitative Research 239

3. Explanatory Nonexperimental Quantitative Research


240

1. Summary 244

2. Key Terms and Concepts 245

3. Related Internet Sites 245

4. Practice Test 245

5. Challenge Exercises 246

2. 13 Survey Research 247

1. Purposes of Survey Research 249

2. Steps in Survey Research 250

3. Cross-Sectional and Longitudinal Designs 251

4. Selecting a Survey Data Collection Method 252

5. Constructing and Refining a Survey Instrument 254

1. Principle 1. Write Items to Match the Research


Objectives 255
2. Principle 2. Write Items That Are Appropriate and
Meaningful for the Respondents to Be Surveyed 255

3. Principle 3. Write Short, Simple Questions 256

4. Principle 4. Avoid Loaded and Leading Questions 256

5. Principle 5. Avoid Double-Barreled Questions 256

6. Principle 6. Avoid Double Negatives 257

7. Principle 7. Determine Whether Closed-Ended and/or


Open-Ended Questions Are Needed 257

8. Principle 8. Construct Mutually Exclusive and


Exhaustive Response Categories for Closed-
Ended Questions 258

9. Principle 9. Consider the Different Types of Closed-


Ended Response Categories Rating Scales 258

1. Binary Forced Choice 260

2. Rankings 260

3. Checklists 261

10. Principle 10. Use Multiple Items to Measure Complex


or Abstract Constructs 261

1. Semantic Differential 261

2. Likert Scaling 262

11. Principle 11. Make Sure the Questionnaire Is Easy to


Use from the Beginning to the End 263

1. Ordering of Questions 263


2. Contingency Questions 263

3. Questionnaire Length 264

4. Response Bias 264

12. Principle 12. Pilot Test the Questionnaire Until It Is


Perfected 265

6. Selecting Your Survey Sample from the Population 265

7. Preparing and Analyzing Your Survey Data 267

1. Summary 268

2. Key Terms and Concepts 268

3. Related Internet Sites 268

4. Practice Test 269

5. Challenge Exercises 269

3. 14 Qualitative and Mixed Methods Research 270

1. Major Characteristics of Qualitative Research 272

2. Research Validity in Qualitative Research 273

1. Descriptive Validity 274

2. Interpretive or Emic Validity 274

3. Theoretical Validity 274

4. Internal (Causation) Validity 274

5. External (Generalizing) Validity 275

3. Four Major Qualitative Research Methods 276


1. Phenomenology 276

1. Phenomenological Data Collection and Data


Analysis 277

2. Phenomenological Report Writing 278

2. Ethnography 279

1. Ethnographic Data Collection Methods 280

2. Entry, Group Acceptance, and Fieldwork 280

3. Data Analysis and Report Writing 281

3. Case Study Research 282

1. Data Collection in Case Study Research 282

2. Case Study Designs 282

3. Case Study Data Analysis and Report Writing 283

4. Grounded Theory 283

1. Data Collection in Grounded Theory Research


285

2. Grounded Theory Data Analysis and Report


Writing 285

4. Mixed Methods Research 286

5. Research Validity in Mixed Methods Research 287

6. Mixed Methods Designs 289

1. Summary 292
2. Key Terms and Concepts 293

3. Related Internet Sites 293

4. Practice Test 294

5. Challenge Exercises 294

6. Part VI Analyzing and Interpreting Data 295

1. 15 Descriptive Statistics 295

1. Descriptive Statistics 296

2. Frequency Distributions 298

3. Graphic Representations of Data 299

1. Bar Graphs 299

2. Histograms 300

3. Line Graphs 300

4. Scatterplots 302

4. Measures of Central Tendency 304

1. Mode 304

2. Median 304

3. Mean 304

5. Measures of Variability 305

1. Range 306

2. Variance and Standard Deviation 306


3. Standard Deviation and the Normal Curve 306

1. z scores 307

6. Examining Relationships Among Variables 309

1. Unstandardized and Standardized Difference


Between Group Means 309

2. Correlation Coefficient 310

1. Partial Correlation Coefficient 315

3. Regression Analysis 315

4. Contingency Tables 318

1. Summary 320

2. Key Terms and Concepts 320

3. Related Internet Sites 320

4. Practice Test 321

5. Challenge Exercises 321

2. 16 Inferential Statistics 322

1. Sampling Distributions 323

2. Estimation 325

3. Hypothesis Testing 327

1. Directional Alternative Hypotheses 331

2. Review of the Logic of Hypothesis Testing 332

3. Hypothesis-Testing Errors 332


4. Hypothesis Testing in Practice 334

1. The t Test for Correlation Coefficients 335

2. One-Way Analysis of Variance 336

3. Post Hoc Tests in Analysis of Variance 337

4. Analysis of Covariance 338

5. Two-Way Analysis of Variance 340

6. One-Way Repeated Measures Analysis of Variance 342

7. The t Test for Regression Coefficients 344

5. Nonparametric Statistics 346

1. Chi-Square Test for Independence 347

2. Other Significance Tests 348

6. Hypothesis Testing and Research Design 348

1. Summary 350

2. Key Terms and Concepts 351

3. Related Internet Sites 352

4. Practice Test 352

5. Challenge Exercises 352

7. Part VII Writing the Research Report 354

1. 17 Preparing the Research Report Using APA Style 354

1. The Apa Format 356


2. Preparation of the Research Report 363

1. Writing Style 364

2. Language 365

1. Specificity 365

2. Labels 365

3. Participation 366

4. Specific Issues 366

3. Editorial Style 367

1. Italics 367

2. Abbreviations 367

3. Headings 367

4. Quotations 368

5. Numbers 368

6. Physical Measurements 368

7. Presentation of Statistical Results 368

8. Tables 368

9. Figures 369

10. Figure Legends and Captions 370

11. Figure Preparation 370

12. Reference Citations 370


13. Reference List 371

14. Preparation of the Manuscript for Submission 372

15. Ordering of Manuscript Pages 372

3. Submission of the Research Report for Publication 373

1. Acceptance of the Manuscript 374

4. Presenting Research Results at Professional Conferences


374

1. Oral Presentation 375

2. Poster Presentation 375

1. Summary 377

2. Key Terms and Concepts 377

3. Related Internet Sites 378

4. Practice Test 378

5. Challenge Exercises 378

1. Appendix 379

2. Glossary 380

3. References 392

4. Index 401
Preface
Welcome to Research Methods, Design, and Analysis. You are embarking
on a study that will help you to think systematically, critically, and
creatively in Psychology and other disciplines. We have two primary goals
for this text. First, we have focused on writing a book that provides an
understanding of the research methods used to investigate human thought
and behavior. Research methods tend to change slowly, but they do change.
This book provides coverage of the complete range of research methods
available today. Psychology tends to favor experimental methods, so we
devote more time to experimental research methods. Because
nonexperimental research also is used in many areas of psychology, we
carefully cover this method, including how to write a proper questionnaire.
Because of the rapid growth of qualitative and mixed methods in
psychology, we carefully cover these methods to complement the more
traditional methods and to add to each student’s repertoire of research
skills.

A second overarching goal that has been maintained throughout all


editions of the textbook is to present information in a way that is
understandable to students. We have attempted to meet this goal by
presenting material in as simple and straightforward a manner as possible
and by accompanying complex material with illustrations taken from the
research literature. We believe that such illustrations not only assist in
clarifying the presented material, but also bring the material to life when it
is placed in the context of actual research studies. This allows the student
not only to learn the material, but also to see how it is used in a research
study.

Overview and Organization


of the Textbook
Research Methods, Design, and Analysis is written at the undergraduate
level and is intended for use in the undergraduate methods course. The
book provides an introduction to all aspects of research methodology, and
assumes no prior knowledge. The chapters are divided into seven major
parts, as follows:
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quite impervious, but becoming—at any rate in the case of the larger
and more important pair—open previous to the final ecdysis. We
have mentioned the contradictory opinions of Réaumur and Dufour,
and will now add the views of some modern investigators. Oustalet
says[341] that there are two pairs of spiracles in the nymphs; the first
pair is quite visible to the naked eye, and is situate between pro- and
meso-notum; it is in the nymph closed by a membrane. The other
pair of spiracles is placed above the posterior pair of legs, is small
and completely closed. He does not state what stage of growth was
attained by the nymphs he examined. Palmén was of opinion that
not only thoracic but abdominal spiracles exist in the nymph,[342] and
that they are completely closed so that no air enters them; he says
that the spiracles have tracheae connected with them, that at each
moult the part closing the spiracles is shed with some of the tracheal
exuviae attached to it. The breathing orifices are therefore for a short
time at each ecdysis open, being subsequently again closed by
some exudation or secretion. This view of Palmén's has been
thought improbable by Hagen and Dewitz, who operated by placing
nymphs in alcohol or warm water and observing the escape of
bubbles from the spots where the supposed breathing orifices are
situate. Both these observers found much difference in the results
obtained in the cases of young and of old nymphs. Hagen concludes
that the first pair of thoracic spiracles are functionally active, and that
abdominal stigmata exist though functionless; he appears to be of
opinion that when the first thoracic stigma is closed this is the result
of the abutting against it of a closed trachea. Dewitz found[343] that
in the adult nymph of Aeschna the thoracic stigma is well developed,
while the other stigmata—to what number and in what position is not
stated—are very small. In a half-grown Aeschnid nymph he found
the thoracic stigma to be present in an undeveloped form. On
placing a full-grown nymph in alcohol, gas escaped from the stigma
in question, but in immature nymphs no escape of gas occurred
although they were subjected to a severe test. A specimen that,
when submitted to the above-mentioned immersion, emitted gas,
subsequently moulted, and thereafter air escaped from the spiracle
previously impervious. The observations of Hagen and Dewitz are
perhaps not so adverse to the views of Palmén as has been
supposed, so that it would not be a matter for surprise if Palmén's
views on this point should be shown to be quite correct.

The number of species of Odonata or Libellulidae that have been


described is somewhat less than two thousand, but constant
additions are made to the number, and when the smaller and more
fragile forms from the tropics are collected and worked out it will
probably be found that the number of existing species is somewhere
between five and ten thousand. They are distributed all over the
world, but are most numerous in species in the warmer regions, and
their predominance in any one locality is very much regulated by the
existence of waters suitable for the early stages of their lives.

A good work on the British Odonata is still a desideratum.[344] In


Britain about forty-six species are believed to be native. They are
said to be of late years less numerous than they used to be.
Notwithstanding their great powers of flight, dragon-flies are
destroyed by birds of various kinds; several hawks are said to be
very fond of them, and Merops persicus to line its nest with their
wings. The number of Insects killed by dragon-flies in places where
they are abundant must be enormous; the nymphs, too, are very
destructive in the waters they inhabit, so that dragon-flies have no
doubt been no mean factor in maintaining that important and delicate
balance of life which it is so difficult for us to appreciate. The nymphs
are no doubt cannibals, and this may perhaps be an advantage to
the species, as the eggs are sometimes deposited in large numbers
in a limited body of water, where all must perish if the nymphs did
not, after exhausting other food, attack one another. Martin, speaking
of the Odonata of the Département de l'Indre in France, says:[345]
"The eggs, larvae, and nymphs are the prey of several fishes,
snakes, newts, Coleoptera, aquatic Hemiptera, and of some diving
birds. Sometimes the destruction is on a considerable scale, and one
may notice the dragon-flies of some piece of water to diminish
gradually in numbers, while the animals that prey on them increase,
so that a species may for a time entirely disappear in a particular
spot, owing to the attacks of some enemy that has been specially
prosperous, and also eager in their pursuit. De Selys found that from
a pond filled with carp, roach, perch, and eels, several of the dragon-
fly denizens disappeared directly the bream was introduced." On the
other hand, there can be little doubt that the nymphs are sometimes
injurious to fish; it has been recorded that in a piscicultural
establishment in Hungary 50,000 young fishes were put into a pond
in spring; in the following autumn only fifty-four fish could be found,
but there were present an enormous quantity of dragon-fly nymphs.

Odonata are among the few kinds of Insects that are known to form
swarms and migrate. Swarms of this kind have been frequently
observed in Europe and in North America; they usually consist of
species of the genus Libellula, but species of various other genera
also swarm, and sometimes a swarm may consist of more than one
species. L. quadrimaculata is the species that perhaps most
frequently forms these swarms in Europe; a large migration of this
species is said to occur every year in the Charente inférieure from
north to south.[346] It is needless to say that the instincts and stimuli
connected with these migrations are not understood.

The nymphs are capable, under certain circumstances, of


accommodating themselves to very peculiar conditions of life. The
Sandwich Islands are extremely poor in stagnant waters, and yet
there exist in this remote archipelago several highly peculiar species
of Agrioninae. Mr. R. C. L. Perkins has recently discovered that the
nymphs of some of these are capable of maintaining their existence
and completing their development in the small collections of water
that accumulate in the leaves of some lilies growing on dry land.
These nymphs (Fig. 271) have a shorter mask than occurs, we
believe, in any other Odonata, and one would suppose that they
must frequently wait long for a meal, as they must be dependent on
stray Insects becoming immersed in these tiny reservoirs. The
cannibal habits of the Odonata probably stand these lily-dwellers in
good stead; Mr. Perkins found that there were sometimes two or
three nymphs of different sizes together, and we may suspect that it
sometimes goes hard with the smaller fry. The extension in the
length of the body of one of these lily-frequenting Agrions when it
leaves the water for its aerial existence is truly extraordinary.

Fig. 271.—Under side of Agrionid nymph, with short mask, living in


water in lilies. Hawaiian Islands. × 3.

The Odonata have no close relations with any other group of Insects.
They were associated by Latreille with the Ephemeridae, in a family
called Subulicornia. The members of the two groups have, in fact, a
certain resemblance in some of the features of their lives, especially
in the sudden change, without intermediate condition, from aquatic to
aerial life; but in all important points of structure, and in their
dispositions, dragon-flies and may-flies are totally dissimilar, and
there is no intermediate group to connect them. We have already,
said that the Odonata consist of two very distinct divisions—
Anisopterides and Zygopterides. The former group comprises the
subfamilies Gomphinae, Cordulegasterinae, Aeschninae,
Corduliinae, and Libellulinae,—Insects having the hinder wings
slightly larger than the anterior pair; while the Zygopterides consist of
only two subfamilies—Calepteryginae and Agrioninae; they have the
wings of the two pairs equal in size, or the hinder a little the smaller.
The two groups Gomphinae and Calepteryginae are each, in several
respects, of lower development than the others, and authorities are
divided in opinion as to which of the two should be considered the
more primitive. It is therefore of much interest to find that there exists
an Insect that shares the characters of the two primitive subfamilies
in a striking manner. This Insect, Palaeophlebia superstes (Fig. 272),
has recently been discovered in Japan, and is perhaps the most
interesting dragon-fly yet obtained. De Selys Longchamps refers it to
the subfamily Calepteryginae, on account of the nature of its wings;
were the Insect, however, deprived of these organs, no one would
think of referring Palaeophlebia to the group in question, for it has
the form, colour, and appearance of a Gomphine Odonate.
Moreover, the two sexes differ in an important character,—the form
of the head and eyes. In this respect the female resembles a
Gomphine of inferior development; while the male, by the shape and
large size of the ocular organs, may be considered to combine the
characters of Gomphinae and Calepteryginae. The Insect is very
remarkable in colour, the large eyes being red in the dead examples.
We do not, however, know what may be their colour during life, as
only one pair of the species is known, and there is no record as to
the life-history and habits. De Selys considers the nearest ally of this
Insect to be Heterophlebia dislocata, a fossil dragon-fly found in the
Lower Lias of England.

Fig. 272.—Palaeophlebia superstes. A, The Insect with wings of one


side and with two legs removed; B, front view of head of female;
C, of male. (After De Selys.)

Numerous fossil dragon-flies are known; the group is well


represented in the Tertiary strata, and specimens have been found in
amber. In strata of the Secondary age these Insects have been
found as far back as the Lower Lias; their remains are said to exist in
considerable variety in the strata of that epoch, and some of them to
testify to the existence at that period of dragon-flies as highly
specialised as those now living. According to Hagen[347]
Platephemera antiqua and Gerephemera simplex, two Devonian
fossils, may be considered as dragon-flies; the evidence as to this
appears inadequate, and Brongniart refers the latter Insect to the
family Platypterides, and considers Platephemera to be more allied
to the may-flies.
One of the most remarkable of the numerous discoveries lately
made in fossil entomology is the finding of remains of huge Insects,
evidently allied to dragon-flies, in the Carboniferous strata at
Commentry. Brongniart calls these Insects Protodonates,[348] and
looks on them as the precursors of our Odonata. Meganeura monyi
was the largest of these Insects, and measured over two feet across
the expanded wings. If M. Brongniart be correct in his restoration of
this giant of the Insect world, it much resembled our existing dragon-
flies, but had a simple structure of the thoracic segments, and a
simpler system of wing-nervures. On p. 276 we figured
Titanophasma fayoli, considered by Scudder and Brongniart as allied
to the family Phasmidae, and we pointed out that this supposed
alliance must at best have been very remote. This view is now taken
by M. Brongniart himself,[349] he having removed the Insect from the
Protophasmides to locate it in the Protodonates near Meganeura.
There appears to be some doubt whether the wings supposed to
belong to this specimen were really such, or belonged rather to
some other species.

CHAPTER XIX

AMPHIBIOUS NEUROPTERA CONTINUED—EPHEMERIDAE, MAY-FLIES

Fam. VII. Ephemeridae—May-flies.

Delicate Insects with atrophied mouth and small, short antennae;


with four membranous wings having much minute cross-veining;
the hinder pair very much smaller than the other pair, sometimes
entirely absent: the body terminated by three or two very
elongate slender tails. The earlier stages are passed through in
water, and the individual then differs greatly in appearance from
the winged Insect; the passage between the two forms is
sudden; the creature in its first winged state is a subimago,
which by shedding a delicate skin reveals the final form of the
individual.

Fig. 273.—Ephemera danica, male, Britain.

The may-flies are well known—in literature—as the types of a brief


and ineffective life. This supposed brevity relates solely to their
existence in the winged form. In the earlier stages the may-fly is so
unlike its subsequent self that it is not recognised as a may-fly by the
uninitiated. The total life of the individual is really quite as long as
that of most other Insects. The earlier stages and life-histories of
these Insects are of great importance. The perfect Insects are so
delicate and fragile that they shrivel much in drying, and are very
difficult to preserve in a condition suitable for study.

The mouth of the imago is atrophied, the trophi scarcely existing as


separate parts. Packard says that in Palingenia bilineata he could
discover no certain traces of any of the mouth-parts, but in
Leptophlebia cupida he found, as he thought, the rudiments of the
maxillae and labium, though not of the mandibles. The antennae are
always short, and consist of one or two thick basal joints succeeded
by a delicate needle-like segment, which, though comparatively long,
is not divided. The ocular organs are remarkable for their large size
and complex development; they are always larger in the male than
they are in the female. The compound eyes of the former sex are in
certain species, e.g. Cloëon (Fig. 274), quite divided, so that each
eye becomes a pair of organs of a different character; one part forms
a pillar facetted at its summit, while the other part remains as a true
eye placed on the side of the head; in front of these compound eyes
there are three ocelli. Thus the Insect comes to have three different
kinds of eyes, together seven in number.

Fig. 274.—Front of head of Cloëon, male. a, Pillared eye; b, sessile


eye; c, ocellus.

The prothorax is small, the pronotum being, however, quite distinct.


The mesothorax is very large; its notum forms by far the larger part
of the upper surface of the thoracic region, the metathorax being
small and different in structure, resembling in appearance a part of
the abdomen, so that the hind wings look as if they were attached to
a first abdominal segment. The mesosternum is also
disproportionately large in comparison with the homologous piece
preceding it, and with that following it. The pleural pieces are large,
but their structure and disposition are only very imperfectly
understood. The coxae are small and are widely separated, the
anterior being, however, more elongate and approximate than the
others. The other parts of the legs are slender; the number of joints
in the tarsi varies from five to one. The legs throughout the family
exhibit a considerable variety of structure, and the front pair in the
males of some species are remarkably long. The abdomen is usually
slender, and consists of ten segments; the terminal one bears three,
or two, very long flexible appendages. The first dorsal plate of the
abdomen is either wanting or is concealed to a considerable extent
by the metanotum. The wings are peculiar; the anterior pair vary a
great deal in their width, but are never very long in proportion to the
width; the hind pair are always disproportionately small, and
sometimes are quite wanting. The venation consists of a few, or of a
moderate number, of delicate longitudinal veins that do not pursue a
tortuous course, but frequently are gracefully curved, and form a
system of approximately similar curves, most of the veins being of
considerable length; close to the anterior margin of the wing there
are two or three sub-parallel veins. Frequently there are very
numerous fine, short cross-veinlets, but these vary greatly and may
be entirely wanting.

Fig. 275.—Wings of Ephemera danica. (After Eaton.)

The earlier stages of the life of Ephemeridae are, it is believed, in the


case of all the species, aquatic. May-flies, indeed, during the period
of their post-embryonic development are more modified for an
aquatic life than any other Insects, and are provided with a complex
apparatus of tracheal gills. The eggs are committed to the waters
without any care or foresight on the part of the parent flies, thus the
embryonic development is also aquatic; little, however, is known of it.
According to Joly[350] the process in Palingenia virgo is slow. The
larva on emerging from the egg has no respiratory system, neither
could Joly detect any circulation or any nervous system. The
creature on emergence is very like Campodea in form, possessing
long antennae and tails—caudal setae. Owing to the organisation
being inferior, the creature in its earlier stages is called a larvule; in
its later stages it is usually spoken of as a nymph, but the term larva
is also frequently applied to it. Soon the gills begin to appear in the
form of small tubular caeca placed in the posterior and upper angles
of the abdominal rings; in fifteen days the gills begin to assume their
characteristic form, are penetrated by tracheae, and the circulation
can be seen. The amount of growth accomplished after hatching
between March and September is but small.
Fig. 276.—Nymph of Cloëon dipterum.[351] Wing-sheath of left side,
gills of right side, removed; g, tracheal gills. (After Vayssière.)

Fig. 277.—Larvule of Cloëon dimidiatum. (After Lubbock.)

The metamorphosis of Cloëon has been described by Sir John


Lubbock; he informs us that the young creature undergoes a
constant and progressive development, going through a series of
more than twenty moults, each accompanied by a slight change of
form or structure. His observations were made on captured
specimens, so that it is not certain that what he calls[352] the first
stage is really such. He found no tracheae in the earliest stages; the
small first rudiments of the gills became visible in the third stage,
when there were no tracheae; the fourth instar possessed tracheae,
and they could be seen in the gills. The wing rudiments could first be
detected in the ninth and tenth stages. The changes of skin during
the winter months are separated by longer intervals than those
occurring at other periods of the year.
Fig. 278.—Adult nymph of Ephemera vulgata. (After Eaton.) Britain.

The nymphs differ greatly in the structure and arrangement of their


tracheal gills, and display much variety in their general form and
habits; some of them are very curious creatures. Pictet[353] divides
them in accordance with their habits into four groups: (1) Fossorial
larvae: these live in the banks of streams and excavate burrows for
shelter; they are of cylindrical form, possess robust legs, abundant
gills at the sides of the body, and frequently processes projecting
forwards from the head: examples, Ephemera (Fig. 278) and
Palingenia. (2) Flat larvae: these live attached to rocks, but run with
rapidity when disturbed; they prefer rapid streams, have the
breathing organs attached to the sides of the body and not reposing
on the back; they are exclusively carnivorous, while the fossorial
forms are believed to obtain their nutriment by eating mud: example,
Baëtis. (3) Swimming larvae: elongate delicate creatures, with feeble
legs, and with strongly ciliated caudal setae: example, Cloëon (Fig.
276). (4) Climbing larvae: these live in slowly-moving waters,
especially such as have much slimy mud in suspension, and they
have a habit of covering themselves with this mud sometimes to
such an extent as to become concealed by it: example,
Potamanthus.
Fig. 279.—Nymph of Oligoneuria garumnica, France. g2 and g7, two of
the dorsal tracheal gills. (After Vayssière.)

The anatomy of the nymphs has been treated by Vayssière,[354] who


arranges them in five groups in accordance with the conditions of the
tracheal gills: (1) The gills are of large size, are exposed and
furnished at the sides with respiratory fringes: example, Ephemera
(Fig. 278). (2) The branchiae are blade-like, not fringed, and are
exposed at the sides of the body: example, Cloëon (Fig. 276). (3)
The respiratory tubes are placed on the under surface of plates
whose upper surface is not respiratory: example, Oligoneuria
garumnica (Fig. 279). (4) The anterior gill is modified to form a plate
that covers the others: example, Tricorythus (Fig. 282, B). (5) The
gills are concealed in a respiratory chamber: example, Prosopistoma
(Fig. 280). The last of these nymphs is more completely adapted for
an aquatic life than any other Insect at present known; it was for long
supposed to be a Crustacean, but it has now been shown to be the
early stage of a may-fly, the sub-imago having been reared from the
nymph. The carapace by which the larger part of the body is covered
is formed by the union of the pro- and meso-thorax with the sheaths
of the anterior wings, which have an unusually extensive
development; under the carapace there is a respiratory chamber, the
floor and sides of which are formed by the posterior wing-sheaths,
and by a large plate composed of the united nota of the metathorax
and the first six abdominal segments. In this chamber there are
placed five pairs of tracheal gills; entrance of water to the chamber is
effected by two laterally-placed orifices, and exit by a single dorsal
aperture. These nymphs use the body as a sucker, and so adhere
strongly to stones under water. When detached they swim rapidly by
means of their caudal setae; the form of these latter organs is
different from that of other Ephemerid nymphs. This point and other
details of the anatomy of this creature have been described in detail
by Vayssière.[355] These nymphs have a very highly developed
tracheal system; they live in rapid watercourses attached to stones
at a depth of three to six inches or more under the water. Species of
Prosopistoma occur in Europe, Madagascar, and West Africa.

Fig. 280.—Prosopistoma punctifrons, nymph. France. (After Vayssière.)


o, Orifice of exit from respiratory chamber.

According to Eaton,[356] in the nymphs of some Ephemeridae the


rectum serves, to a certain extent, as a respiratory agent; he
considers that water is admitted to it and expelled after the manner
we have described in Odonata, p. 421.

Fig. 281.—A, Last three abdominal segments and bases of the three
caudal processes of Cloëon dipterum: r, dorsal vessel; kl, ostia
thereof; k, special terminal chamber of the dorsal vessel with its
entrance a; b, blood-vessel of the left caudal process; B, twenty-
sixth joint of the left caudal process from below; b, a portion of the
blood-vessel; o, orifice in the latter. (After Zimmermann.)

The internal anatomy of the nymphs of Ephemeridae shows some


points of extreme interest. The long caudal setae are respiratory
organs of a kind that is almost if not quite without parallel in the other
divisions of Insecta. The dorsal vessel for the circulation of the blood
is elongate, and its chambers are arranged one to each segment of
the body. It drives the blood forwards in the usual manner, but the
posterior chamber possesses three blood-vessels, one of which is
prolonged into each caudal seta. This terminal chamber is so
arranged as to drive the blood backwards into the vessels of the
setae; on the under surface of the vessels there are oval orifices by
which the blood escapes into the cavity of the seta so as to be
submitted to the action of the surrounding medium for some of the
purposes of respiration. This structure has been described by
Zimmermann,[357] who agrees with Creutzberg[358] that the organ by
which the blood is propelled into the setae is a terminal chamber of
the dorsal vessel; Verlooren,[359] who first observed this accessory
system of circulation, thought the contractile chamber was quite
separate from the heart. The nature of the connexion between this
terminal chamber that drives the blood backwards and the other
chambers that propel the fluid forwards appears still to want
elucidation.

Fig. 282.—A, Nymph of Ephemerella ignita with gills of left side


removed; g, gills: B, nymph of Tricorythus sp. with gill cover of
right side removed; g.c, gill cover; g, g′, gills. (After Vayssière.)

The nymphs of the Ephemeridae being creatures adapted for


existence in water, the details of their transformation into creatures
having an entirely aerial existence cannot but be of much interest. In
the nymphs the tracheal system is well developed, but differs from
that of air-breathing Insects in the total absence of any spiracles.
Palmén has investigated this subject,[360] and finds that the main
longitudinal tracheal trunks of the body of the nymph are not
connected with the skin of the body by tracheae, but are attached
thereto by ten pairs of slender strings extending between the
chitinous integument and the tracheal trunks. When the skin is shed
these strings—or rather a chitinous axis in each one—are drawn out
of the body, and bring with them the chitinous linings of the tracheae.
Thus notwithstanding the absence of spiracles, the body wall is at
each moult pierced by openings that extend to the tracheae. After
the ordinary moults these orifices close immediately, but at the
change to the winged state they remain open and form the spiracles.
At the same time the tracheal gills are completely shed, and the
creature is thus transformed from a water-breather to an Insect
breathing air as usual. In addition to this change there are others of
great importance, such as the development of the great eyes and the
complete atrophy of the mouth-parts. The precise manner of these
changes is not known; they occur, however, within the nymph skin.
The sudden emergence of the winged Insect from the nymph is one
of the most remarkable facts in the life-history of the may-fly; it has
been observed by Sir John Lubbock,[361] who describes it as almost
instantaneous. The nymph floats on the water, the skin of the back
opens, and the winged Insect flies out, upwards and away; "from the
moment when the skin first cracks not ten seconds are over before
the Insect has flown away." The creature that thus escapes has not,
however, quite completed its transformation. It is still enveloped in a
skin that compresses and embarrasses it; this it therefore rapidly
gets rid of, and thus becomes the imago, or final instar of the life-
cycle. The instar in which the creature exists winged and active,
though covered with a skin, is called the sub-imago. The parts of the
body in the sub-imago are as a whole smaller than they are in the
imago, and the colour is more dingy; the appendages—wings, legs,
and caudal setae—are generally considerably shorter than they are
in the imago, but attain their full length during the process of
extraction. The creatures being, according to Riley, very impatient
and eager to take to the wing, the completion of the shedding of the
skin of the sub-imago is sometimes performed while the Insect is
flying in the air.

Fig. 283.—Lingua of Heptagenia longicauda, × 16. m, Central; l, lateral


pieces. (After Vayssière.)

The food of young Ephemeridae is apparently of a varied and mixed


nature. Eaton says[362] that though sometimes the stronger larvae
devour the weaker, yet the diet is even in these cases partly
vegetable. The alimentary canal frequently contains much mud; very
small organisms, such as diatoms and confervae, are thought to
form a large part of the bill of fare of Ephemerid nymphs. Although
the mouth is atrophied in the imago, yet it is highly developed in the
nymphs. This is especially notable in the case of the lingua or
hypopharynx (Fig. 283); indeed Vayssière[363] seems to incline to the
opinion that this part of the mouth may be looked on in these Insects
as a pair of appendages of a head-segment (see p. 96 ante), like the
labium or maxillae.

The life-history has not been fully ascertained in the case of any
species of may-fly; it is known, however, that the development of the
nymph sometimes occupies a considerable period, and it is thought
that in the case of some species this extends to as much as three
years. It is rare to find the post-embryonic development of an Insect
occupying so long a period, so that we are justified in saying that
brief as may be the life of the may-fly itself, the period of preparation
for it is longer than usual. Réaumur says, speaking of the winged fly,
that its life is so short that some species never see the sun. Their
emergence from the nymph-skin taking place at sunset, the duties of
the generation have been, so far as these individuals are concerned,
completed before the morning, and they die before sunrise. He
thinks, indeed, that individuals living thus long are to be looked on as
Methuselahs among their fellows, most of whom, he says, live only
an hour or half an hour.[364] It is by no means clear to which species
these remarks of Réaumur refer; they are doubtless correct in
certain cases, but in others the life of the adult is not so very short,
and in some species may, in all probability, extend over three or four
days; indeed, if the weather undergo an unfavourable change so as
to keep them motionless, the life of the flies may be prolonged for a
fortnight.

The life of the imago of the may-fly is as remarkable as it is brief; in


order to comprehend it we must refer to certain peculiarities of the
anatomy with which the vital phenomena are connected. The more
important of these are the large eyes of the males, the structure of
the alimentary canal, and that of the reproductive organs. We have
already remarked that the parts of the mouth in the imago are
atrophied, yet the canal itself not only exists but is even of greater
capacity than usual; it appears to have much the same general
arrangement of parts as it had in the nymph. Its coats are, however,
of great tenuity, and according to Palmén[365] the divisions of the
canal are separated by changes in the direction of certain portions
anterior to, and of others posterior to, its central and greater part—
the stomach—in such a manner that the portions with diverted
positions act as valves. The stomach, in fact, forms in the interior of
the body a delicate capacious sac; when movement tends to
increase the capacity of the body cavity then air enters into the
stomachic sac by the mouth orifice, but when muscular contractions
result in pressure on the sac they close the orifices of its extremities
by the valve-like structures we have mentioned above; the result is,
that as complex movements of the body are made the stomach
becomes more and more distended by air. It was known even to the
old naturalists that the dancing may-fly is a sort of balloon, but they
were not acquainted with the exact mode of inflation. Palmén says
that in addition to the valve-like arrangements we have described,
the entry to the canal is controlled by a circular muscle, with which
are connected radiating muscles attached to the walls of the head.
Palmén's views are adopted, and to a certain extent confirmed, by
Fritze,[366] who has examined the alimentary canal of the may-fly,
and considers that though the normal parts of the canal exist, the
function is changed in the imago, in which the canal serves as a sort
of balloon, and aids the function of the reproductive organs. The
change in the canal takes place in an anticipatory manner during the
nymph and sub-imago stages.

The sexual organs of Ephemeridae are remarkable for their


simplicity; they are destitute of the accessory glands and diverticula
that, in some form or other, are present in most other Insects. Still
more remarkable is the fact that the ducts by which they
communicate with the exterior continue as a pair to the extremity of
the body, and do not, as in other Insects, unite into a common duct.
Thus in the female there is neither bursa copulatrix, receptaculum
seminis, nor uterine portion of oviduct, and there is no trace of an
ovipositor; the terminations of the ducts are placed at the hind
margin of the seventh ventral plate, just in front of which they are
connected by a fold of the integument. The ovary consists of a very
large number of small egg-tubes seated on one side of a sac, which
forms their calyx, and one of whose extremities is continued
backwards as one of the pair of oviducts. The male has neither
vesiculae seminales, accessory glands, nor ductus ejaculatorius.
The testes are elongate sacs, whose extremities are prolonged
backwards forming the vasa deferentia; these open separately at the
extremity of the body, each on a separate intromittent projection of
more or less complex character, the two organs being, however,
connected by means of the ninth ventral plate, of which they are,
according to Palmén, appendages. We should remark that this
authority considers Heptagenia to form, to some extent, an exception
as regards the structures of the female; while Polymitarcys is in the
male sex strongly aberrant, as the two vasa deferentia, instead of
being approximately straight, are bent inwards at right angles near
their extremities so as to meet, and form in the middle a common
cavity, which then again becomes double to pass into the pair of
intromittent organs.
According to the views of Exner and others, the compound eyes of
Insects are chiefly organs for the perception of movement; if this
view be correct, movements such as those made during the dances
of may-flies may, by the number of the separate eyes, by their
curved surfaces and innumerable facets, be multiplied and
correlated in a manner of which our own sense of sight allows us to
form no conception. We can see on a summer's evening how
beautifully and gracefully a crowd of may-flies dance, and we may
well believe that to the marvellous ocular organs of the flies
themselves (Fig. 274) these movements form a veritable ballet. We
have pointed out that by this dancing the peculiarly formed
alimentary canal becomes distended, and may now add that Palmén
and Fritze believe that the unique structure of the reproductive
organs is also correlated with the other anatomical peculiarities, the
contents of the sexual glands being driven along the simple and
direct ducts by the expansion of the balloon-like stomach. During
these dances the momentary conjugation of the sexes occurs, and
immediately thereafter the female, according to Eaton, resorts to the
waters appropriate for the deposition of her eggs. As regards this,
Eaton says:[367] "Some short-lived species discharge the contents of
their ovaries completely en masse, and the pair of fusiform or
subcylindrical egg-clusters laid upon the water rapidly disintegrate,
so as to let the eggs sink broadcast upon the river-bed. The less
perishable species extrude their eggs gradually, part at a time, and
deposit them in one or other of the following manners: either the
mother alights upon the water at intervals to wash off the eggs that
have issued from the mouths of the oviducts during her flight, or else
she creeps down into the water to lay her eggs upon the under-side
of stones, disposing them in rounded patches, in a single layer
evenly spread, and in mutual contiguity." The eggs are very
numerous, and it is thought may sometimes remain in the water as
much as six or seven months before they hatch.

The number of individuals produced by some kinds of may-flies is


remarkable. Swarms consisting of millions of individuals are
occasionally witnessed. D'Albertis observed Palingenia papuana in
countless myriads on the Fly River in New Guinea: "For miles the
surface of the river, from side to side, was white with them as they
hung over it on gauzy wings; at certain moments, obeying some
mysterious signal, they would rise in the air, and then sink down
anew like a fall of snow." He further states that the two sexes were in
very disproportionate numbers, and estimates that there was but a
single female to every five or six thousand males.

Ephemeridae in the perfect state are a favourite food of fishes, and it


is said that on some waters it is useless for the fly-fisher to try any
other lure when these flies are swarming. Most of the "duns" and
"spinners" of the angler are Ephemeridae; so are several of the
"drakes," our large E. danica and E. vulgata being known as the
green drake and the gray drake. Ronalds says[368] that the term
"dun" refers to the pseud-imago condition, "spinner" to the perfect
Insect. E. danica and E. vulgata are perhaps not distinguished by
fishers; Eaton says that the former is abundant in rapid, cool
streams, while E. vulgata prefers warmer and more tranquil rivers.

These sensitive creatures are unable to resist the attractions of


artificial lights. Réaumur noticed this fact many years ago, and since
the introduction of the electric light, notes may frequently be seen in
journals recording that myriads of these Insects have been lured by it
to destruction. Their dances may frequently be observed to take
place in peculiar states of light and shade, in twilight, or where the
sinking sun has its light rendered broken by bushes or trees;
possibly the broken lights are enhanced in effect by the ocular
structures of the Insects. It has recently been ascertained that a
species of Teleganodes is itself luminous. Mr. Lewis,[369] who
observed this Insect in Ceylon, states that in life the whole of the
abdomen was luminous, not brightly so, but sufficient to serve as a
guide for capturing the Insect on a dark night. It has also been
recorded that the male of Caenis dimidiata gives a faint blue light at
night.

Nearly 300 species of Ephemeridae are known, but this may be only
a fragment of what actually exist, very little being known of may-flies

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