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2019v1.0
SIXTH EDITION

CLINICAL
HEMATOLOGY
ATLAS
Jacqueline H. Carr, MS MT(ASCP)SH
Laboratory Manager (Retired)
Department of Pathology and Laboratory Medicine
Indiana University Health
Indianapolis, Indiana
Elsevier
3251 Riverport Lane
St. Louis, Missouri 63043

CLINICAL HEMATOLOGY ATLAS, SIXTH EDITION ISBN: 978-0-323-71192-0

Copyright © 2022 by Elsevier, Inc. All rights reserved

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Notice

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Printed in Canada

Last digit is the print number: 9 8 7 6 5 4 3 2 1

 Dedicated to my dear friend and colleague,
Bernadette Rodak,
who ended her battle with cancer on March 22, 2016.
Also
My husband,
Charles Carr,
who has supported me through this adventure
as well as our daughters,
Kimberly Mayrose and Alexis Carr.
REVIEWERS

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PREFACE

B
ecause the emphasis of an atlas is morphology, the Clinical Hematology Atlas
is intended to be used with a textbook, such as Rodak’s Hematology, sixth edi-
tion, that addresses physiology and diagnosis along with morphology. This atlas
is designed for a diverse audience that includes clinical laboratory science students,
medical students, residents, and practitioners. It is also a valuable resource for clinical
laboratory practitioners who are being retrained or cross-trained in hematology. It is
not intended to be a detailed, comprehensive manual for diagnosis.
In this concise format, every photomicrograph and word has been evaluated
for value to the microscopist. All superuous information has been excluded in an
attempt to maintain focus on signicant microscopic ndings while correlating this
information with clinical diagnosis. What started as a primer for Clinical Laboratory
Science students with no previous hematology education has evolved into an inter-
nationally recognized reference for multiple levels of expertise, from entry level to
practicing professionals.

ORGANIZATION

As is frequently expounded, morphology on a peripheral blood lm is only as good as

the quality of the smear and the stain. Chapter 1 reviews smear preparation, staining,
and the appropriate area in which to evaluate cell distribution and morphology. A
table that summarizes the morphology of leukocytes found in a normal differential,
along with multiple examples of each cell type, facilitates early instruction in blood
smear review.
Chapter 2 schematically presents hematopoietic features of cell maturation.
General cell maturation, along with an electron micrograph with labeled organelles,
will help readers correlate the substructures with the appearance of cells under light
microscopy. Visualizing normal cellular maturation is essential to the understanding
of disease processes. This correlation of schematic, electron micrograph, and Wright-
stained morphology is carried throughout the maturation chapters. Figure 2-1 has
been formatted to reect recent hematopoietic theory. In addition, the chart aids
readers in recognizing the anatomical sites at which each stage of maturation nor-
mally occurs.
Chapters 3 to 9 present the maturation of each cell line individually, repeating
the respective segment of the overall hematopoietic scheme from Chapter 2, to
assist the student in seeing the relationship of each cell line to the whole. In these
chapters, each maturation stage is presented as a color print, a schematic, and an
electron micrograph. A description of each cell, including overall size, nuclear-to-
cytoplasmic ratio, morphologic features, and reference ranges in peripheral blood and
bone marrow, serves as a convenient summary. The nal gure in each of these chap-
ters summarizes lineage maturation by repeating the hematopoietic segment with
the corresponding photomicrographs. Multiple nomenclatures for erythrocyte mat-
uration are used to accommodate use in multiple settings and demographic groups.

viii
 Preface ix

Chapters 10 to 12 present discrete cellular abnormalities of erythrocytes, that is,

variations in size, color, shape, and distribution, as well as inclusions found in eryth-
rocytes. Each variation is presented along with a description of the abnormality, or
composition of the inclusion, and associated disorders.
Because diseases are often combinations of the cellular alterations, Chapter 13
integrates morphologic ndings into the diagnostic features of disorders primarily
affecting erythrocytes.
In Chapter 14, nuclear and cytoplasmic changes in leukocytes are displayed and
correlated with non-malignant leukocyte disorders.
Diseases of excessive or altered production of cells may be caused by maturation
arrest, asynchronous development, or proliferation of one cell line, as presented in
Chapters 15 to 19. Cytochemical stains are presented with disorders in which they
are useful.
The therapeutic use of myeloid growth factors causes morphologic changes that
mimic severe infections or malignancies. Chapter 20 presents examples of peripheral
blood morphology following G-CSF or GM-CSF. It is the authors’ design that the cel-
lular defects in leukocyte disorders be visually compared with the process of normal
hematopoiesis for a more thorough comprehension of normal and altered develop-
ment. Readers are encouraged to refer to the normal hematopoiesis illustration, Figure
2-1, for comparison of normal and abnormal cells and the progression of diseases.
Microorganisms, including parasites, may be seen on peripheral blood smears.
A brief photographic overview is given in Chapter 21. Readers are encouraged to
consult a microbiology reference, such as Mahon CM, Lehman DC, Manuselis G:
Textbook of Diagnostic Microbiology, fth edition, for a more detailed presentation.
Chapter 22 includes photomicrographs that are not categorized into any one
particular area, such as fat cells, mitotic gures, metastatic tumor cells, and artifacts.
Chapter 23 describes ndings expected in the peripheral blood of neonates, includ-
ing anticipated variations in morphology and cellular distribution. Comparison of the
hematogone, normal for newborns, with the blast cell of acute leukemia is included.
Chapter 24 is intended to be an overview of the most frequent microscopic nd-
ings in body uids. It is not proposed as a comprehensive review of the cytology of
human body uids, but rather a quick reference for the beginning microscopist as
well as the seasoned professional.
As with previous editions, the sixth edition features spiral binding, making the
atlas more convenient when used at the microscope bench.
All of these chapters combine into what we believe is a comprehensive and valu-
able resource for any clinical laboratory. The quality of the schematic illustrations,
electron micrographs, and color photographs stand for themselves. We hope that
this atlas will enrich the learning process for the student and serve as an important
reference tool for the practitioner.

EVOLVE

The Evolve website provides free materials for both students and instructors.
Instructors have access to an electronic image collection featuring all of the images
from the atlas. Students and instructors have access to student review questions and
summary tables.
ACKNOWLEDGMENTS

A
s in previous edition, the faculty and staff of the Department of Pathology
and Laboratory Medicine, Indiana University School of Medicine have been
supportive off this effort. An incredibly special thanks is owed to George
Girgis who willingly and unselshly shared his vast knowledge of microscopy in
blood cell and body uid morphology. Without his help and support, this edition
of the Clinical Hematology Atlas would not have been possible. A special thanks to
Linda Marler and Jean Siders who once again allowed us to publish their microbi-
ology images.
The professionals at Elsevier who been incredibly patient during the last year as
this atlas began to come to life. The following people deserve my gratitude for their
kind assistance and advice:
Heather Bays-Petrovic, Content Strategist
Abigail Bradberry, Senior Project Manager
Maria Broeker, Senior Content Development Specialist

x
CONTENTS

Section 1 Introduction
1 Introduction to Peripheral Blood Film Examination 1

Section 2 Hematopoiesis
2 Hematopoiesis 11
3 Erythrocyte Maturation 17
4 Megakaryocyte Maturation 31
5 Neutrophil Maturation 41
6 Eosinophil Maturation 55
7 Basophil Maturation 65
8 Monocyte Maturation 69
9 Lymphocyte Maturation 79

Section 3 Erythrocytes
10 Variations in Size and Color of Erythrocytes 89
11 Variations in Shape and Distribution of Erythrocytes 93
12 Inclusions in Erythrocytes 107
13 Diseases Affecting Erythrocytes 115

Section 4 Leukocytes
14 Nuclear and Cytoplasmic Changes in Leukocytes 131
15 Acute Myeloid Leukemia 141
16 Precursor Lymphoid Neoplasms 161
17 Myeloproliferative Neoplasms 165
18 Myelodysplastic Syndromes 175
19 Mature Lymphoproliferative Disorders 185
20 Morphologic Changes After Myeloid Hematopoietic Growth Factors 195

Section 5 Miscellaneous
21 Microorganisms 199
22 Miscellaneous Cells 207
23 Normal Newborn Peripheral Blood Morphology 219
24 Body Fluids 223

Glossary 245
Index 265

xi
This page intentionally left blank
CHAPTER 1

INTRODUCTION TO PERIPHERAL
BLOOD FILM EXAMINATION
2 SECTION 1 Introduction

A
properly prepared blood lm is essential to accurate assessment of cellular
morphology. A variety of methods are available for preparing and staining
blood lms, the most common of which are discussed in this atlas. It is beyond
the scope of this atlas to discuss other methodologies; however, detailed descrip-
tions of these procedures can be found in textbooks on hematology, such as Rodak’s
Hematology: Clinical Principles and Applications, sixth edition.

WEDGE FILM PREPARATION

MAKING THE PERIPHERAL BLOOD FILM

Although some automated analyzers prepare and stain blood lms according to
established criteria, manual blood lm preparation is still used in many places. The
wedge lm is a convenient and commonly used technique for making peripheral
blood lms. This technique requires at least two 3 × 1-inch (75 × 25-mm) clean
glass slides. High-quality, beveled-edge microscope slides are recommended. One
slide serves as the blood lm slide, and the other as the spreader slide. These can
then be reversed to prepare a second lm. A drop of ethylenediaminetetraacetic
acid (EDTA) anticoagulated blood about 3 mm in diameter is placed at one end
of the slide. Alternatively, a similar size drop of blood directly from a nger or heel
puncture is acceptable. The size of the drop of blood is important. Too large a drop
creates a long or thick lm, and too small a drop often makes a short or thin lm.
In preparing the lm, the technician holds the pusher slide securely in front of the
drop of blood at a 30- to 45-degree angle to the lm slide (Figure 1.1A). The pusher
slide is pulled back into the drop of blood and held in that position until the blood
spreads across the width of the slide (Figure 1.1B). It is then quickly and smoothly
pushed forward to the end of the lm slide, creating a wedge lm (Figure 1.1C).
It is important that the whole drop of blood is picked up and spread. Moving the
pusher slide forward too slowly accentuates poor leukocyte distribution by pushing
larger cells, such as monocytes and granulocytes, to the very ends and sides of the
lm. Maintaining a consistent angle between the slides and an even, gentle pressure
is essential. It is frequently necessary to adjust the angle between the slides to pro-
duce a satisfactory lm. For higher than normal hematocrit, the angle between the
slides must be lowered so that the lm is not too short and thick. For extremely low
hematocrit, the angle must be raised. A well-made peripheral blood lm (Figure 1.2)
has the following characteristics:
1. About two-thirds to three-fourths of the length of the slide is covered by the lm.
2. It is slightly rounded at the feather edge (thin portion), not bullet shaped.
3. Lateral edges of the lm should be visible. The use of slides with chamfered (bev-
eled) corners may facilitate this appearance.
4. It is smooth without irregularities, holes, or streaks.
5. When the slide is held up to light, the feather edge of the lm should have a
“rainbow” appearance.
6. The whole drop is picked up and spread.
Figure 1.3 shows examples of unacceptable lms.
 CHAPTER 1 Introduction to Peripheral Blood Film Examination 3

30 – 45°

C
FIGURE 1.1 Wedge technique of making a peripheral blood lm. (A) Correct angle to hold spreader slide.
(B) Blood spread across width of slide. (C) Completed wedge lm.
(From Keohane E.A., Smith L., Walenga J. (Eds.) (2016). Rodak’s hematology: clinical principles and
applications. (5th ed.). St. Louis: Saunders Elsevier.)

FIGURE 1.2 Well-made peripheral

blood lm.
(From Keohane E.A., Smith L., Walenga
J. (Eds.) (2016). Rodak’s hematology:
clinical principles and applications. (5th
ed.). St. Louis: Saunders Elsevier.)
4 SECTION 1 Introduction

A B C D

E F G H
FIGURE 1.3 Unacceptable peripheral blood lms. Slide appearances associated with the most com-
mon errors are shown, but note that a combination of causes may be responsible for unacceptable
lms. (A) Chipped or rough edge on spreader slide. (B) Hesitation in forward motion of spreader slide.
(C) Spreader slide pushed too quickly. (D) Drop of blood too small. (E) Drop of blood not allowed to
spread across the width of the slide. (F) Dirt or grease on the slide; may also be caused by elevated
lipids in the blood specimen. (G) Uneven pressure on the spreader slide. (H) Time delay; drop of blood
began to dry.
(From Keohane E.A., Smith L., Walenga J. (Eds.) (2016). Rodak’s hematology: clinical principles and
applications. (5th ed.). St. Louis: Saunders Elsevier.)
 CHAPTER 1 Introduction to Peripheral Blood Film Examination 5

STAINING OF PERIPHERAL BLOOD FILMS

The purpose of staining blood lms is to identify cells and recognize morphology
easily through the microscope. Wright or Wright-Giemsa stains are the most com-
monly used for peripheral blood and bone marrow lms. These stains contain both
eosin and methylene blue and are therefore termed polychrome stains. The colors
vary slightly from laboratory to laboratory, depending on the method of staining.
Slides must be allowed to dry thoroughly before staining. The cells are xed to
the glass slide by the methanol in the stain. Staining reactions are pH dependent,
and the actual staining of the cellular components occurs when a buffer (pH 6.4)
is added to the stain. Free methylene blue is basic and stains acidic cellular compo-
nents, such as RNA, blue. Free eosin is acidic and stains basic components, such as
hemoglobin or eosinophilic granules, red. Neutrophils have cytoplasmic granules
that have a neutral pH and accept some characteristics from both stains. Details
for specic methods of staining peripheral blood and bone marrow lms, including
automated methods, may be found in a standard textbook of hematology.
An optimally stained lm (Figure 1.4) has the following characteristics:
1. The red blood cells (RBCs) should be pink to salmon.
2. Nuclei are dark blue to purple.
3. Cytoplasmic granules of neutrophils are lavender to lilac.
4. Cytoplasmic granules of basophils are dark blue to black.
5. Cytoplasmic granules of eosinophils are red to orange.
6. The area between the cells should be colorless, clean, and free of precipitated stain.
A well-stained slide is necessary for accurate interpretation of cellular morphology.
The best staining results are obtained from freshly made slides that have been pre-
pared within 2 to 3 hours of blood collection. Box 1.1 lists common reasons for
poorly stained slides and may be used as a guide when troubleshooting.

FIGURE 1.4 Optimally stained peripheral blood

lm demonstrating the appropriate area in
which to perform the white blood cell differ-
ential and morphology assessment and the
platelet estimate. Only the center of the eld is
shown; an entire eld would contain 200 to 250
red blood cells (original × 1000).
6 SECTION 1 Introduction

Box 1.1
Troubleshooting Poorly Stained Blood Films
First Scenario
Problems
• Red blood cells appear gray
• White blood cells are too dark
• Eosinophil granules are gray, not orange

Causes
• Stain or buffer too alkaline (most common)
• Inadequate rinsing
• Prolonged staining
• Heparinized blood sample

Second Scenario
Problems
• Red blood cells too pale or red color
• White blood cells barely visible

Causes
• Stain or buffer too acidic (most common)
• Underbuffering (time too short)
• Over-rinsing

From Keohane E.A., Smith L., Walenga J. (Eds.) (2016). Rodak’s

hematology: clinical principles and applications. (5th ed.).
St. Louis: Saunders Elsevier.

PERIPHERAL FILM EXAMINATION

10 × EXAMINATION
Examination of the blood lm is a multistep process. Begin the lm examination
with a scan of the slide using the 10 × or low-power objective (total magnication
= 100 ×). This step is necessary to assess the overall quality of the lm, including
abnormal distribution of RBCs, suggesting the presence of rouleaux or autoagglu-
tination, and/or the presence of a disproportionate number of large nucleated cells
such as monocytes or neutrophils at the edges of the lm. If the latter exists, another
lm should be prepared. In addition, the 10 × lm examination allows for the rapid
detection of large abnormal cells such as blasts, reactive lymphocytes, and parasites.

40 × OR 50 × EXAMINATION
Using the 40 × (high dry) objective or the 50 × oil objective (400 × and 500 × total
magnication, respectively), nd an area of the lm in which the RBCs are evenly dis-
tributed and barely touching one another (two or three cells may overlap; Figure 1.5).
Scan 8 to 10 elds in this area of the lm, and determine the average number of
white blood cells (WBCs) per eld. Although an exact factor varies with the make
Another random document with
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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