A Text-book of Entomology - Including the Anatomy, Physiology, Embryology and Metamorphoses of Insects for Use in Agricultural and Technical Schools and Colleges As Well As by the Working Entomologist

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Title: A Text-book of Entomology - Including the Anatomy, Physiology, Embryology and Metamorphoses of Insects for Use in Agricultural and Technical Schools and Colleges As Well As by the Working Entomologist
Author: Packard, Alpheus S.
Language: English
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A TEXT-BOOK OF ENTOMOLOGY

[Illustration]

A
TEXT-BOOK OF ENTOMOLOGY
INCLUDING
THE ANATOMY, PHYSIOLOGY, EMBRYOLOGY AND METAMORPHOSES
OF
INSECTS
_FOR USE IN AGRICULTURAL AND TECHNICAL SCHOOLS AND COLLEGES_
AS WELL AS BY THE WORKING ENTOMOLOGIST

ALPHEUS S. PACKARD, M.D., PH.D.

PROFESSOR OF ZOÖLOGY AND GEOLOGY, BROWN UNIVERSITY AUTHOR OF “GUIDE TO
STUDY OF INSECTS,” “ENTOMOLOGY FOR BEGINNERS,” ETC.

New York
THE MACMILLAN COMPANY
LONDON: MACMILLAN & CO., LTD.
1898

_All rights reserved_

Norwood Press
J. S. Cushing & Co.—Berwick & Smith
Norwood Mass. U.S.A.

PREFACE

[Illustration]

In preparing this book the author had in mind the wants both of the
student and the teacher. For the student’s use the more difficult
portions, particularly that on the embryology, may be omitted. The work
has grown in part out of the writer’s experience in class work.

In instructing small classes in the anatomy and metamorphoses of
insects, it was strongly felt that the mere dissection and drawing of a
few types, comprising some of our common insects, were by no means
sufficient for broad, thorough work. Plainly enough the laboratory work
is all important, being rigidly disciplinary in its methods, and
affording the foundation for any farther work. But to this should be
added frequent explanations or formal lectures, and the student should
be required to do collateral reading in some general work on structural
and developmental entomology. With this aim in view, the present work
has been prepared.

It might be said in explanation of the plan of this book, that the
students having previously taken a lecture course in the zoölogy of the
invertebrates, were first instructed in the facts and conclusions
bearing on the relations of insects to other Arthropoda, and more
especially the anatomy of Peripatus, of the Myriopoda, and of
Scolopendrella. Then the structure of Campodea, Machilis, and Lepisma
was described, after which a few types of winged insects, beginning with
the locust and ending with the bee, were drawn and dissected; the nymph
of the locust, and the larva and pupa of a moth and of a wasp and bee
being drawn and examined. Had time permitted, an outline of the
embryology and of the internal changes in flies during their
metamorphoses would have been added.

This book gives, of course with much greater fulness and detail for
reference and collateral reading, what we roughly outlined in our class
work. The aim has been to afford a broad foundation for future more
special work by any one who may want to carry on the study of some group
of insects, or to extend in any special direction our present knowledge
of insect morphology and growth.

Many of our entomologists begin their studies without any previous
knowledge of the structure of animals, taking it up as an amusement.
They may be mere collectors and satisfied simply to know the name of
their captures, but it is hoped that with this book in their hands they
may be led to desire farther information regarding what has already been
done on the structure and mode of growth of the common insects. For
practical details as to how to dissect, to make microscopic slides, and
to mount and preserve insects generally, they are referred to the
author’s “Entomology for Beginners.”

It may also be acknowledged that even in our best and latest general
treatises on zoölogy, or comparative anatomy, or morphology, the portion
related to insects is scarcely so thoroughly done as those parts devoted
to other phyla, that of Lang, however, his invaluable Comparative
Anatomy, being an exception. On this account, therefore, it is hoped
that this hiatus in our literature may be in a degree filled.

The author has made free use of the excellent article “Insecta” of
Newport, of Lang’s comprehensive summary in his most useful Text-book of
Comparative Anatomy, of Graber’s excellent Die Insecten, of Miall and
Denny’s The Structure and Life-History of the Cockroach, and of Sharp’s
Insecta. Kolbe’s Einführung has been most helpful. But besides these
helps, liberal use has been made of the very numerous memoirs and
monographic articles which adorn our entomological literature. The
account of the embryology of insects is based on Korschelt and Heider’s
elaborate work, Lehrbuch der Vergleichenden Entwicklungsgeschichte der
Wirbellosen Thiere, the illustrations of this portion being mainly taken
from it, through the Messrs. Swan Sonnenschein & Co., London.

Professor H. S. Pratt has kindly read over the manuscript and also the
proofs of the portion on embryology and metamorphoses, and the author is
happy to acknowledge the essential service he has rendered.

The bibliographical lists are arranged by dates, so as to give an idea
of the historical development of each subject. The aim has been to make
these lists tolerably complete and to include the earliest, almost
forgotten works and articles as well as the most recent.

Much care has been taken to give due credit either to the original
sources from which the illustrations are copied, or to the artist; about
ninety of the simpler figures were drawn by the author, many of them for
this work. For the use of certain figures acknowledgments are due to the
Boston Society of Natural History, to the Division of Entomology, U. S.
Department of Agriculture, through the kind offices of Mr. L. O. Howard,
and to the Illinois State Laboratory of Natural History, through
Professor S. A. Forbes and Mr. C. A. Hart. Professor W. M. Wheeler, of
the University of Chicago, has kindly loaned for reproduction several of
his original drawings published in the Journal of Morphology. A number
are reproduced from figures in the reports of the United States
Entomological Commission.

PROVIDENCE, R. I.,
March 4, 1898.

TABLE OF CONTENTS

[Illustration]

PART I. MORPHOLOGY AND PHYSIOLOGY

PAGE
POSITION OF INSECTS IN THE ANIMAL KINGDOM 1

RELATIONS OF INSECTS TO OTHER ARTHROPODA 2

The Crustacea 4
The Merostomata 5
The Trilobita 5
The Arachnida 6
Relations of Peripatus to insects 9
Relation of Myriopods to insects 11
Relations of the Symphyla to insects 18
Diagnostic or essential characters of Symphyla 22

INSECTA (HEXAPODA) 26

Diagnostic characters of insects 26

1. EXTERNAL ANATOMY

_a._ Regions of the body 27

_b._ The integument (exoskeleton) 28

Chitin 29

_c._ Mechanical origin and structure of the segments (somites,
arthromeres, etc.) 30

_d._ Mechanical origin of the limbs and of their jointed
structure 35

THE HEAD AND ITS APPENDAGES 42

_a._ The head 42

The labrum 42
The epipharynx and labrum-epipharynx 43
Attachment of the head to the trunk 46
The basal or gular region of the head 46
The occiput 48
The tentorium 49
Number of segments in the head 50
The composition of the head in the Hymenoptera 55

_b._ Appendages of the head 57

The antennæ 57
The mandibles 59
The first maxillæ 62
The second maxillæ 68
The hypopharynx 70
Does the hypopharynx represent a distinct segment? 82

THE THORAX AND ITS APPENDAGES 86

_a._ The thorax: its external anatomy 86

The patagia 89
The tegulæ 89
The apodemes 92
The acetabula 94

_b._ The legs: their structure and functions 95

Tenent hairs 99
Why do insects have but six legs? 100
Loss of limbs by disuse 101

_c._ Locomotion (walking, climbing, and swimming) 103

Mechanics of walking 103
Locomotion on smooth surfaces 111
Climbing 116
The mode of swimming of insects 116

_d._ The wings and their structure 120

The veins 121
The squamæ 123
The halteres 124
The thyridium 124
The tegmina and hemelytra 124
The elytra 124

_e._ Development and mode of origin of the wings 126

Embryonic development of the wings 126
Evagination of the wing outside of the body 132
Extension of the wing; drawing out of the tracheoles 133

_f._ The primitive origin of the wings 137

The development and structure of the tracheæ and veins of
the wing 144

_g._ Mechanism of flight 148

Theory of insect flight 150
Graber’s views as to the mechanism of the wings, flight,
etc. 153

THE ABDOMEN AND ITS APPENDAGES 162

The median segment 163
The cercopoda 164
The ovipositor and sting 167
The styles and genital claspers (Rhabdopoda) 176
Velum penis 181
The suranal plate 181
The podical plates or paranal lobes 182
The infra-anal lobe 183
The egg-guide 183

THE ARMATURE OF INSECTS: SETÆ, HAIRS, SCALES, TUBERCLES, ETC. 187

The cuticula 187
Setæ 188
Glandular hairs and spines 190
Scales 193
Development of the scales 195
Spinules, hair-scales, hair-fields, and androconia 197

THE COLORS OF INSECTS 201

Optical colors 201
Natural colors 203
Chemical and physical nature of the pigment 206
Ontogenetic and phylogenetic development of colors 207

2. INTERNAL ANATOMY

THE MUSCULAR SYSTEM 211

Musculature of a caterpillar 213
Musculature of a beetle 213
Minute structure of the muscles 215
Muscular power of insects 217

THE NERVOUS SYSTEM 222

_a._ The nervous system as a whole 222

_b._ The brain 226

The optic or procerebral segment 231
Procerebral lobes 232
The mushroom or stalked bodies 233
Structure of the mushroom bodies 234
The central body 237
The antennal or olfactory lobes (Deutocerebrum) 237
The œsophageal lobes (Tritocerebrum) 237

_c._ Histological elements of the brain 238

_d._ The visceral (sympathetic or stomatogastric) system 238

_e._ The supraspinal cord 240

_f._ Modifications of the brain in different orders of insects 240

_g._ Functions of the nerve-centres and nerves 243

THE SENSORY ORGANS 249

_a._ The eyes and insect vision 249

The simple or single-lensed eye (ocellus) 249
The compound or facetted eye (ommateum) 250
The facet or cornea 250
The crystalline lens or cone 251
The pigment 253
The basilar membrane 253
The optic tract 253
Origin of the facetted eye 255
Mode of vision by single eyes or ocelli 255
Mode of vision by facetted eyes 256
The principal use of the facetted eye to perceive the
movements of animals 259
How far can insects see? 260
Relation of sight to the color of eyes 260
The color sense of insects 260

_b._ The organs of smell 264

Historical sketch of our knowledge of the organs of smell 264
Physiological experiments 268
Relation of insects to smelling substances before and
after the loss of their antennæ 269
Experiments on the use of the antennæ in seeking for food 270
Experiments testing the influence of the antennæ of the
males in seeking the females 270
Structure of the organs of smell in insects 271

_c._ The organs of taste 281

Structure of the taste organs 282
Distribution in different orders of insects 282
Experimental proof 286

_d._ The organs of hearing 287

The ears or tympanal and chordotonal sense-organs of
Orthoptera and other insects 288
Antennal auditory hairs 292
Special sense-organs in the wings and halteres 293

_e._ The sounds of insects 293

THE DIGESTIVE CANAL AND ITS APPENDAGES 297

_a._ The digestive canal 302

The œsophagus 303
The crop or ingluvies 303
The “sucking stomach” or food-reservoir 305
The fore-stomach or proventriculus 306
The œsophageal valve 311
Proventricular valvule 313
The peritrophic membrane 313
The mid-intestine 314
Histology of the mid-intestine 316
The hind-intestine 316
Large intestine 316
The ileum 317
The gastro-ileal folds 317
The colon 317
The rectum 318
The vent (anus) 319
Histology of the digestive canal 320

_b._ Digestion in insects 324

The mechanism of secretion 326
Absorbent cells 328

THE GLANDULAR AND EXCRETORY APPENDAGES OF THE DIGESTIVE CANAL 331

_a._ The salivary glands 331

_b._ The silk or spinning glands, and the spinning apparatus 339

The process of spinning 340
How the thread is drawn out 343
Appendages of the silk-gland (Filippi’s glands) 345

_c._ The cæcal appendages 347

_d._ The excretory system (urinary or Malpighian tubes) 348

Primitive number of tubes 353

_e._ Poison-glands 357

_f._ Adhesive or cement-glands 360

_g._ The wax-glands 361

_h._ “Honey-dew” or wax-glands of Aphids 364

_i._ Dermal glands in general 365

DEFENSIVE OR REPUGNATORIAL SCENT-GLANDS 368

Eversible coxal glands 369
Fœtid glands of Orthoptera 369
Anal glands of beetles 372
The blood as a repellent fluid 374
Eversible glands of caddis-worms and caterpillars 375
The osmeterium in Papilio larvæ 377
Dorsal and lateral eversible metameric sacs in other larvæ 377
Distribution of repugnatorial or alluring scent-glands in
insects 382

THE ALLURING OR SCENT-GLANDS 391

THE ORGANS OF CIRCULATION 397

_a._ The heart 397

The propulsatory apparatus 401
The supraspinal vessel 403
The aorta 404
The pericardial cells 405
Pulsatile organs of the legs 405

_b._ The blood 407

The leucocytes 407

_c._ The circulation of the blood 409

Effects of poisons on the pulsations 412

THE BLOOD TISSUE 419

_a._ The fat-body 419

_b._ The pericardial fat-body or pericardial cells 420

Leucocytes or phagocytes in connection with the
pericardial cells 421

_c._ The œnocytes 423

_d._ The phosphorescent organs 424

Physiology of the phosphorescence 426

THE RESPIRATORY SYSTEM 430

_a._ The tracheæ 431

Distribution of the tracheæ 432

_b._ The spiracles or stigmata 437

The position and number of pairs of stigmata 439
The closing apparatus of the stigma 441

_c._ Morphology and homologies of the tracheal system 442

_d._ The spiral threads or tænidia 444

_e._ Origin of the tracheæ and of the “spiral thread” 447

Internal, hair-like bodies 451

_f._ The mechanism of respiration and the respiratory
movements of insects 451

_g._ The air-sacs 456

The use of the air-sacs 457

_h._ The closed or partly closed tracheal system 459

_i._ The rectal, tracheal gills, and rectal respiration of
larval Odonata and other insects 463

_j._ Tracheal gills of the larvæ of insects 466

Blood-gills 475

_k._ Tracheal gills of adult insects 476

THE ORGANS OF REPRODUCTION 485

_a._ The male organs of reproduction 494

The testes 495
The seminal ducts 496
The ejaculatory duct 497
The accessory glands 497
The spermatozoa 497
Formation of the spermatozoön 498

_b._ The female organs of reproduction 500

The ovaries and the ovarian tubes 500
Origin of incipient eggs in the germ of the testes 504
The bursa copulatrix 505
The spermatheca 506
The colleterial glands 506
The vagina or uterus 507
Signs of copulation in insects 507

PART II. EMBRYOLOGY OF INSECTS

_a._ The egg 515

Mode of deposition 518
Vitality of eggs 520
Appearance and structure of the ripe egg 520
The egg-shell and yolk-membrane 520
The micropyle 522
Internal structure of the egg 524

_b._ Maturation or ripening of the egg 525

_c._ Fertilization of the egg 525

_d._ Division and formation of the blastoderm 526

_e._ Formation of the first rudiments of the embryo and of the
embryonic membranes 531

Formation of the embryonic membranes 532
The gastrula stage 535
Division of the embryo or primitive band into
body-segments 536
Differences between the invaginated and overgrown
primitive band 538
Revolution of the embryo where the primitive band is
invaginated 540

_f._ Formation of the external form of the body 542

Origin of the body-segments 542
The procephalic lobes 544
Fore-intestine (stomodæum) and hind-intestine
(proctodæum), labrum 547
Completion of the head 548

_g._ The appendages 548

The cephalic appendages 548
The thoracic appendages 550
The abdominal appendages 550
Appendages of the first abdominal segment (pleuropodia) 551
Are the abdominal legs of Lepidoptera and phytophagous
Hymenoptera true limbs? 552
The tracheæ 553

_h._ Nervous system 554

Completion of the definite form of the body 555

_i._ Dorsal closure and involution of the embryonic membranes 556

_j._ Formation of the germ-layers 558

_k._ Farther development of the mesoderm; formation of the
body-cavity 563

_l._ Formation of organs 566

The nervous system 566
Development of the brain 567
Development of the eyes 567
Intestinal canal and glands 569
The salivary glands 570
The urinary tubes 572
The heart 572
The blood-corpuscles 574
Musculature; connective tissue; fat-body 574
The reproductive organs 575
Development of the male germinal glands 579

_m._ Length of embryonic life 582

_n._ The process of hatching 583

The hatching spines 585

PART III. THE METAMORPHOSES OF INSECTS

_a._ The nymph as distinguished from the larval stage 593

_b._ Stages or stadia of metamorphosis 594

_c._ Ametabolous and metabolous stages 594

THE LARVA 599

_a._ The Campodea-form type of larva 600

_b._ The eruciform type of larva 602

_c._ Growth and increase in size of the larva 608

_d._ The process of moulting 609

The number of moults in insects of different orders 615
Reproduction of lost limbs 619
Formation of the cocoon 619
Sanitary conditions observed by the honey-bee larva, and
admission of air within the cocoon 623

THE PUPA STATE 625

_a._ The pupa considered in reference to its adaptation to its
surroundings and its relation to phylogeny 631

_b._ Mode of escape of the pupa from its cocoon 632

_c._ The cremaster 636

Mode of formation of the cremaster and suspension of the
chrysalis in butterflies 637

FORMATION OF THE PUPA AND IMAGO IN THE HOLOMETABOLOUS INSECTS (THE
DIPTERA EXCEPTED) 640

_a._ The Lepidoptera 642

The changes in the head and mouth-parts 646
The change in the internal organs 647
The wings 654
Development of the feet and of the cephalic appendages 654
Embryonic cells and the phagocytes 655
Formation of the femur and of the tibia; transformation of
the tarsus 656
The antennæ 657
Maxillæ and labial palpi 658
Process of pupation 660

_b._ The Hymenoptera 661

Ocular or oculo-cephalic buds 665
The antennal buds 665
The buds of the buccal appendages 665
The buds of the ovipositor 665

DEVELOPMENT OF THE IMAGO IN THE DIPTERA 666

_a._ Development of the outer body-form 668

Formation of the imago in Corethra 668
Formation of the imago in Culex 670
Formation of the imago in Chironomus 671
Formation of the imago in Muscidæ 673

_b._ Development of the internal organs of the imago 678

The hypodermis 678
The muscles 680
The digestive canal 681
The tracheal system 683
The nervous system 684
The fat-body 685
Definitive fate of the leucocytes 685
The post-embryonic changes and imaginal buds in the
Pupipara (Melophagus) 686

_c._ General summary 687

HYPERMETAMORPHISM 688

SUMMARY OF THE FACTS AND SUGGESTIONS AS TO THE CAUSES OF
METAMORPHISM 705

Theoretical conclusions; causes of metamorphosis 708

TEXT-BOOK OF ENTOMOLOGY

[Illustration]

PART I.—MORPHOLOGY AND PHYSIOLOGY

[Illustration]

POSITION OF INSECTS IN THE ANIMAL KINGDOM

Although the insects form but a single class of the animal kingdom, they
are yet so numerous in orders, families, genera, and species, their
habits and transformations are so full of instruction to the biologist,
and they affect human interests in such a variety of ways, that they
have always attracted more attention from students than any other class
of animals, the number of entomologists greatly surpassing that of
ornithologists, ichthyologists, or the special students of any other
class, while the literature has assumed immense proportions.

Insects form about four-fifths of the animal kingdom. There are about
250,000 species already named and contained in our museums, while the
number of living and fossil species in all is estimated to amount to
between one and two millions.

In their structure insects are perhaps more complicated than any other
animals. This is partly due to the serial arrangement of the segments
and the consequent segmental repetition of organs, especially of the
external appendages, and of the muscles, the tracheæ, and the nerves.
The brain is nearly or quite as complicated as that of the higher
vertebrates, while the sense-organs, especially those of touch, sight,
and smell are, as a rule, far more numerous and only less complex than
those of vertebrates. Moreover, in their psychical development, certain
insects are equal, or even superior, to any other animals, except birds
and mammals.

The animal kingdom is primarily divided into two grand divisions, the
one-celled (_Protozoa_) and many-celled animals (_Metazoa_). In the
latter group the cells and tissues forming the body are arranged in
three fundamental cell-layers; viz. the _ectoderm_ or outer layer, the
_mesoderm_, and _endoderm_. The series of branches, or phyla, comprised
under the term Metazoa are the Porifera, Cœlenterata, Vermes,
Echinodermata, Mollusca, Arthropoda, and Vertebrata. Their approximate
relationships may be provisionally expressed by the following

TABULAR VIEW OF THE EIGHT BRANCHES OR PHYLA OF THE ANIMAL KINGDOM.

RELATIONS OF INSECTS TO OTHER ARTHROPODA

The insects by general consent stand at the head of the Arthropoda.
Their bodies are quite as much complicated or specialized, and indeed,
when we consider the winged forms, more so, than any other class of the
branch, and besides this they have wings, fitting them for an aërial
life. It is with little doubt that to their power of flight, and thus of
escaping the attacks of their creeping arthropod enemies, insects owe,
so to speak, their success in life; _i.e._ their numerical superiority
in individuals, species, and genera. It is also apparently their power
of moving or swimming swiftly from one place to another which has led to
the numerical superiority in species of fishes to other Vertebrata.
Among terrestrial vertebrates, the birds, by virtue of their ability to
fly, greatly surpass in number of species the reptiles and mammals.

The Arthropoda are in general characterized by having the body composed
of segments (somites or arthromeres) bearing jointed appendages. They
differ from the worms in having segmented appendages, _i.e._ antennæ,
jaws, and legs, instead of the soft unjointed outgrowths of the annelid
worms. Moreover, their bodies are composed of a more or less definite
number of segments or rings, grouped either into a head-thorax
(cephalothorax) and hind-body, as in Crustacea, or into a head
differentiated from the rest of the body (trunk), the latter not being
divided into a distinct thorax and abdomen, as in Myriopoda; or into
three usually quite distinct regions—the head, thorax, and hind-body or
abdomen, as in insects. In certain aberrant, modified forms, as the
Tardigrada, or the Pantopoda, and the mites, the body is not
differentiated into such definite regions.

In their internal organs arthropods agree in their general relations
with the higher worms, hence most zoölogists agree that they have
directly originated from the annelid worms.

The position and general shape of the digestive canal, of the nervous
and circulatory systems, are the same in Arthropoda as in annelid
(oligochete) worms, so much so that it is generally thought that the
Arthropoda are the direct descendants of the worms. It is becoming
evident, however, that there was no common ancestor of the Arthropoda as
a whole, and that the group is a polyphyletic one. Hence, though a
convenient group, it is a somewhat artificial one, and may eventually be
dismembered into at least three or four phyla or branches.

The following diagram may serve to show in a tentative way the relations
of the classes of Arthropoda to each other, and also may be regarded as
a provisional genealogical tree of the branch.

We will now rapidly review the leading features of the classes of
Arthropoda.

=The Crustacea.=—These Arthropoda are in many most important
characteristics unlike the insects; they have two pairs of antennæ, five
pairs of buccal appendages, and they are branchiate Arthropoda. They
have evidently originated entirely independently, and by a direct line
of descent from some unknown annelid ancestor which was either a
many-segmented worm, with parapodia, or the two groups together with the
Rotifera may have originated from a common appendigerous Trochosphæra.
Their segments in the higher forms are definite in number (23 or 24) and
arranged into two regions, a head-thorax (cephalothorax) and hind-body
(abdomen). Nearly all the segments, both of the cephalothorax and
abdomen, bear a pair of jointed limbs, and to them at their base are, in
the higher forms, appended the gills (branchiæ). The limbs are in the
more specialized forms (shrimps and crabs) differentiated into
eye-stalks, two pairs of antennæ, a pair of palpus-bearing jaws
(mandibles), two pairs of maxillæ and three pairs of maxillipeds; these
appendages being biramose, and the latter bearing gills attached to
their basal joints. The legs are further differentiated into ambulatory
thoracic legs and into swimming or abdominal legs, and in the latter the
first pair of the male is modified into copulatory organs (gonopoda).
The male and female reproductive organs as a rule are in separate
individuals, hermaphrodites being very unusual, and the glands may be
paired or single. The sexual outlets are generally paired, and, as in
the male lobster and other Macrura, open in the basal joint of the last
pair of legs, and in the female in the third from the last; while
originally in all Crustacea the sexual organs were most probably paired
(Fig. 3, _B_).

They are, except a few land Isopoda, aquatic, mostly marine, and when
they have a metamorphosis, pass through a six-legged larval stage,
called the Nauplius, the shrimps and crabs passing through an additional
stage, the Zoëa. Crustacea also differ much from insects in the highly
modified nature of the nephridia, which are usually represented by the
green gland of the lobster, or the shell-glands of the Phyllopoda, which
open out in one of the head-segments; also in the possession of a pair
of large digestive glands, the so-called liver.

Intermediate in some respects between the Crustacea and insects, but
more primitive, in respect to what are perhaps the most weighty
characters, than the Crustacea, are the Trilobita, the Merostomata
(Limulus), and, finally, the Arachnida, these being allied groups. In
the Trilobita and Merostomata (Limulus), the head-appendages are more
like feet than jaws, while they have in most respects a similar mode of
embryonic development, the larval forms being also similar.

[Illustration:

FIG. 1.—Restoration of under side of a trilobite (_Triarthrus becki_),
the trunk limbs bearing small triangular respiratory lobes or
gills.—After Beecher.
]

=The Merostomata.=—The only living form, Limulus, is undoubtedly a very
primitive type, as the genital glands and ducts are double, opening wide
apart on the basal pair of abdominal legs (Fig. 3). Moreover, their
head-appendages, which are single, with spines on the basal joint, are
very primitive and morphologically nearer in shape to those of the worms
(Syllidæ, etc.) than even those of the Crustacea. Besides, their four
pairs of coxal glands, with an external opening at the base of the fifth
pair of head-appendages, and which probably are modified nephridia
(Crustacea having but a single pair in any one form, either opening out
on the second antennal, green gland, or second maxillary, shell-gland,
segment), indicate a closer approximation to the polynephrous worms.
Limulus has other archaic features, especially as regards the structure
of the simple and compound eyes and the simple nature of the brain.

=The Trilobita.=—These archaic forms are still more generalized and
primitive than the Merostomata and Crustacea, and probably were the
first Arthropoda to be evolved from some unknown annelid worm. They had
jointed biramose limbs of nearly uniform shape and size on each segment
of the body, which were not, as in Crustacea, differentiated into
antennæ, jaws (mandibles), maxillæ, maxillipeds, and two kinds of legs
(thoracic and abdominal), showing that they are a much more primitive
type, and nearer to the annelids than any other Arthropoda. Their gills,
as shown by the researches of Walcott and of Beecher, were attached to
nearly if not every pair of limbs behind the antennæ (Figs. 1, 2). The
fact that in Trilobita the first pair of limbs is antenniform does not
prove that they are Crustacea, since Eurypterus has a similar pair of
appendages.

[Illustration:

FIG. 2.—Restored section of Calymene: _C_, carapace; _en_, endopodite;
_en′_, exopodite; with the gills on the epipodal or respiratory part
of the appendage.—After Walcott.
]

The limbs in trilobites, as well as the abdominal ones of merostomes,
and all those of Crustacea, except the first antennæ, are biramose,
consisting of an outer (exopodite) and an inner division (endopodite).
In this respect the terrestrial air-breathing tracheate forms,
Arachnida, Myriopoda, and Insecta, differ from the branchiate forms, as
their legs are single or undivided, being adapted for supporting the
body during locomotion upon the solid earth. It is to be observed that
when, as in Limulus, the body is supported by cephalic ambulatory limbs,
they are single, while the abdominal limbs, used as they are in
swimming, are biramose, much as in Crustacea.

=The Arachnida.=—The scorpions and spiders are much less closely allied
to the myriopods and insects than formerly supposed. Their embryology
shows that they have descended from forms related to Limulus, possibly
having had an origin in common with that animal, or having, as some
authors claim, directly diverged from some primitive eurypteroid
merostome. But they differ in essential respects, and not only in the
nature and grouping of their appendages; the first pair instead of
antenniform being like mandibles, and the second pair like the maxillæ,
with the palps, of insects, the four succeeding segments (thoracic)
bearing each a pair of legs. They also have a brain quite unlike that of
Limulus, the nervous cord behind the brain, however, being somewhat
similar, though that of Limulus differs in being enveloped by an
arterial coat. Arachnida respire by tracheæ, besides book-lungs, which,
however, are possibly derivatives of the book-gills of Limulus, while
they perform the office of excretion by means of the malpighian tubes,
and like Limulus possess two large digestive glands (“liver”). Their
embryos have, on at least six abdominal segments, rudiments of limbs,
three pairs of which form the spinnerets, showing their origin from
Limulus-like or eurypteroid forms; their coxal glands are retained from
their eurypteroid ancestors. The Arachnida probably descended from
marine merostomes, and not from an independent annelid ancestry, hence
we have represented them in the diagram on p. 3 as branching off from
the merostomatous phylum, rather than from an independent one.

[Illustration:

FIG. 3.—Paired genital openings of different classes of arthropods.
_A_, the most primitive, of _Limulus polyphemus_: _gen. p_,
generative papillæ; _d_, duct; _vd_, vas deferens; _t_, tendinous
stigmata; _stig_, stigmata; _e_, external branchial muscle; _ant_,
anterior lamellar muscle.—After Benham, with a few changes. _B_,
lobster (_Homarus vulgaris_), ♀: _oe_, genital aperture on 3d pair
of legs; _ov_, ovary; _u_, unpaired portion of the same; _od_,
oviduct. _C_, ♀, scorpion: _ov_, ovary, with a single external
opening. _D_, ♂: _t_, testis; _vd_, vasa deferentia; _sb_, seminal
vesicle; _a_, glandular appendage; _p_, penis.—After Blanchard. _E_,
a myriopod (_Glomeris marginata_, ♀): _os_, ovarian sac, laid open;
_od_, paired oviducts. _F_, ♂: _t_, testis; _gvd_, common vas
deferens; _pa_, paired ducts.—After Favre, from Lang. _G_, _Lepisma
saccharina_, young ♂: _vd_, vas deferens, _ed_, ejaculatory duct;
_ga_, external appendages.—After Nassonow. _H_, Ephemera, ♂, showing
the double outlets.—After Palmén.
]

The characters in which arachnids approach insects, such as tracheæ and
malpighian tubes (none occur, as a rule, in marine or branchiate
arthropods), may be comparatively recent structures acquired during a
change from a marine to a terrestrial life, and not primitive heirlooms.

Arachnida also show their later origin than merostomes by the fact that
their sexual glands are in most cases single, and though with rare
exceptions the ducts are paired, these finally unite and open externally
by a common single genital aperture in the median line of the body, at
the base of the abdomen (Fig. 3, _C_, _D_). In this respect Limulus,
with its pair of genital male or female openings, situated each at the
end of a papilla, placed widely apart at the base of the first abdominal
limbs, is decidedly more archaic. Unlike Crustacea and insects,
Arachnida do not, except in the mites (Acarina), which is a very much
modified group, undergo a metamorphosis.

We see, then, that the insects, with the Myriopoda, are somewhat
isolated from the other Arthropoda. The Myriopoda have a single pair of
antennæ, and as they have other characters in common with insects, Lang
has united the two groups in a single class _Antennata_; but, as we
shall see, this seems somewhat premature and unnecessary. Yet the two
groups have perhaps had a common parentage, and may prove to belong to a
distinct, common phylum.

Not only by their structure and embryology, as well as their
metamorphosis, do the myriopods and insects stand apart from the
Arachnida and other arthropods, but it seems probable that they have had
a different ancestry, the arthropods being apparently polyphyletic.

There are two animals which appear to connect the insects with the
worms, and which indicate a separate line of descent from the worms
independent of that of the other classes. These are the singular
Peripatus, which serves as a connecting link between arthropods and
worms, and Scolopendrella (Symphyla). These two animals are guide-posts,
pointing out, though vaguely to be sure, the way probably trod by the
forms, now extinct, which led up to the insects.

=Relations of Peripatus to Insects.=—We will first recount the
characteristics of this monotypic class. Peripatus (Fig. 4) stands
alone, with no forms intermediate between itself and the worms on the
one hand, and the true Arthropoda on the other. Originally supposed to
be a worm, it is now referred to a class by itself, the Malacopoda of
Blainville, or Protracheata of Haeckel. It lives in the tropics, in damp
places under decaying wood. In general appearance it somewhat resembles
a caterpillar, but the head is soft and worm-like, though it bears a
pair of antenna-like tentacles. It may be said rather to superficially
resemble a leech with clawed legs, the skin and its wrinkles being like
those of a leech. There is a pair of horny jaws in the mouth, but these
are more like the pharyngeal teeth of worms than the jaws of arthropods.
The numerous legs end each in a pair of claws. The ladder-like nervous
system is unlike that of annelid worms or arthropods, but rather recalls
that of certain molluscs (Chiton, etc.), as well as that of certain flat
and nemertine worms. Its annelid features are the large number of
segmentally arranged true nephridia, and the nature of the integument.
Its arthropodan features, which appear to take it out of the group of
worms, are the presence of tracheæ, of true salivary and slime glands,
of a pair of coxal glands (Fig. 4, _C_, _cd_) as well as the claws at
the end of the legs. The tracheæ, which are by no means the only
arthropodan features, are evidently modified dermal glands. The heart is
arthropodan, being a dorsal tube lying in a pericardial sinus, with many
openings. This assemblage of characters is not to be found in any marine
or terrestrial worm.

The tracheæ (Fig. 4, _D_, _tr_) are unbranched fine tubes, without a
“spiral thread,” and are arranged in tufts, in _P. edwardsii_ opening by
simple orifices or pores (“stigmata”) scattered irregularly over the
surface of the body; but in another species (_P. capensis_) some of the
stigmata are arranged more definitely in longitudinal rows,—on each side
two, one dorsally and one ventrally. “The stigmata in a longitudinal row
are, however, more numerous than the pairs of legs.” (Lang.)

The salivary glands, opening by a short common duct into the under side
of the mouth, in the same general position as in insects, are evidently,
as the embryology of the animal proves, transformed nephridia, and being
of the arthropodan type explain the origin and morphology of those of
insects. It is so with the slime glands; these, with the coxal glands,
being transformed and very large dermal glands. Those of insects arose
in the same manner, and are evidently their homologues, while those of
Peripatus were probably originally derived from the setiparous glands in
the appendages (parapodia) of annelid worms.

[Illustration:

FIG. 4.—_A_, _Peripatus novæ zealandiæ_.—After Sedgwick, from Lang.
_B_, _Peripatus capensis_, side view, enlarged about twice the
natural size.—After Moseley, from Balfour. _C_, Anatomy of
_Peripatus capensis_. The enteric canal behind the pharynx has been
removed. _g_, brain; _a_, antenna; _op_, oral or slime papillæ;
_sd_, slime gland; _sr_, slime reservoir, which at the same time
acts as a duct to the gland; _so_{4}_, _so_{5}_, _so_{6}_, _so_{9}_,
nephridia of the 4th, 5th, 6th, and 9th pairs of limbs; _cd_,
elongated coxal gland of the last pair of feet; _go_, genital
aperture; _an_, anus; _ph_, pharynx; _n_, longitudinal trunk of the
nervous system.—After Balfour, from Lang. _D_, Portion of the body
of _Peripatus capensis_ opened to show the scattered tufts of
tracheæ (_tr_); _v_, _v_, ventral nerve cords.—After Moseley.
]

The genital glands and ducts are paired, but it is to be observed that
the outlets are single and situated at the end of the body. In the male
the ejaculatory duct is single; in its base a spermatophore is formed.
It will be seen, then, that Peripatus is not only a composite type, and
a connecting link between worms and tracheate arthropods, but that it
may reasonably be regarded, if not itself the ancestor, as resembling
the probable progenitor of myriopods and insects, though of course there
is a very wide gap between Peripatus and the other antennate,
air-breathing Arthropoda.

[Illustration:

FIG. 4.—_E_, _Peripatus edwardsii_, head from the under side: _a_,
base of antenna; _op_, oral papilla; the figure also shows the
papillæ around the mouth, and the four jaws.—After Balfour, from
Lang. _F_, Anterior end of _Peripatus capensis_, ventral side, laid
open: _a_, antenna; _z_, tongue; _k_, jaw; _sd_, salivary gland;
_gs_, union of the two salivary glands; _ph_, pharynx; _œ_,
œsophagus; _l_, lip papillæ around the mouth; _op_, oral or slime
papilla; _sld_, duct or reservoir of the slime gland.—After Balfour,
from Lang.
]

=Relation of Myriopods to Insects.=—The Myriopoda are the nearest allies
of the insects. They have a distinct head, with one pair of antennæ. The
eyes are simple, with the exception of a single genus (Cermatia), in
which they are aggregated or compound. The trunk or body behind the head
is, as a rule, long and slender, and composed of a large but variable
number of segments, of equal size and shape, bearing jointed legs, which
invariably end in a single claw.

The mouth-parts of the myriopods are so different in shape and general
function from those of insects, that this character, together with the
equally segmented nature of the portion of the body behind the head (the
trunk), forbids our merging them, as some have been inclined to do, with
the insects. There are two sub-classes of myriopods, differing in such
important respects that by Pocock[1] and by Kingsley they are regarded
as independent classes, each equivalent to the insects.

Of these the most primitive are the Diplopoda (Chilognatha), represented
by the galley-worms (Julus, etc.).

[Illustration:

FIG. 5.—Mandible of Julus: _l_, lacinia; _g_, galea; _p_, dens
mandibularis; _ma_, “mala”; _lt_, lamina tritoria; _st_, stipes;
_c_, cardo; _m_, muscle.—After Latzel.
]

In the typical Diplopoda the head consists of three segments, a preoral
or antennal, and two postoral, there being two pairs of jaw-like
appendages, which, though in a broad morphological sense homologues of
the mandibles and first maxillæ of insects, are quite unlike them in
details.

[Illustration:

FIG. 6.—Under lip or deutomala of _Scoterpes copei_: _hyp_, hypostoma
or mentum; _lam. lab_, lamina labialis; _stip. e_, stipes exterior;
with the malella exterior (_mal. e_) and malella interior (_mal.
i_); the stipes interior, with the malulella; and the labiella
(hypopharynx of Vom Rath) with its stilus (_stil._).
]

As we have previously stated,[2] the so-called “mandibles” of diplopods
are entirely different from those of insects, since they appear to be 2–
or 3–jointed, the terminal joint being 2–lobed, thus resembling the
maxillæ rather than the mandibles of insects, which consist of but a
single piece or joint, probably the homologue of the galea or molar
joint of the diplopod protomala. The mandible of the Julidæ (Fig. 5,
_Julus molybdinus_), Lysiopetalidæ, and Polydesmidæ consists of three
joints; viz. a basal piece or cardo, a stipes, and the _mala
mandibularis_, which supports two lobes analogous to the galea and
lacinia of the maxilla of an insect. There is an approach, as we shall
see, in the mandible of Copris, to that of the Julidæ, but in insects in
general the lacinia is wanting, and the jaw consists of but a single
piece.

The deutomalæ (gnathochilarium), or second pair of diplopod jaws, are
analogous to the labium or second maxillæ of insects, forming a
flattened, plate-like under-lip, constituting the floor of the mouth
(Fig. 6). This pair of appendages needs farther study, especially in the
late embryo, before it can be fully understood. So far as known, judging
by Metschnikoff’s work on the embryology of the diplopods, these
myriopods seem to have in the embryo but two pairs of post-antennal
mouth-parts, which he designated as the “mandibles” and “labium.”
Meinert, however, regards as a third pair of mouth-parts or “labium”
what in our Fig. 7 is called the internal stipes (_stip. i._), behind
which is a triangular plate, _lamina labialis_ (_lam. lab_), which he
regards as the sternite of the same segment.

[Illustration:

FIG. 7.—Deutomala of Julus, the lettering as in Fig. 6.
]

[Illustration:

FIG. 8.—Head of _Scolopendra_, seen from beneath, showing the
“mandible” (protomala) with its cardo (_card._) and stipes (_st._),
also the labrum and epilabrum.
]

The hypopharynx, our “labiella,” (Fig. 6), with the supporting rods or
_stili linguales_ (_sti. l_), of Meinert, are of nearly the same shape
as in some insects.

Of the clypeus of insects there is apparently no homologue in myriopods,
though in certain diplopods there is an interantennal clypeal region.
The labium of insects is represented by a short, broad piece, which,
however, unlike that of insects, is immovable, and is flanked by a
separate piece called the _epilabrum_ (Fig. 8). Vom Rath has observed an
epipharynx, which has the same general relations as in insects.

[Illustration:

FIG. 9.—Larva of Julus: _a_, the 3d abdominal segment, with the new
limbs just budding out; _b_, new segments arising between the
penultimate and the last segment.—After Newport.
]

The embryology of myriopods is in many respects like that of insects.
The larva of diplopods hatches with but few segments, and with but three
pairs of limbs; but these are not, as in insects, appended to
consecutive segments, but in one species the third, and in another,
_Julus multistriatus?_ (Fig. 10), the second, segment from the head is
footless, while Vom Rath represents the first segment of an European
Blaniulus as footless, the feet being situated consecutively on segments
2 to 4. The new segments arise at “the growing point” situated between
the last and penultimate segment, growing out in groups of sixes
(Newport) or in our _Julus multistriatus?_ in fives (Fig. 10). In adult
life diplopods (Julus) have a single pair of limbs on the three first
segments, or those corresponding to the thoracic segments of insects,
the succeeding segments having two pairs to each segment.

[Illustration:

FIG. 10.—Freshly hatched larva of _Julus multistriatus?_ 3 mm. long:
_a_, 5 pairs of rudimentary legs, one pair to a segment.
]

Sinclair (Heathcote) regards each double segment in the diplopods as
not two original segments fused together, nor a single segment
bearing two pairs of legs, but as “two complete segments perfect in
all particulars, but united by a large dorsal plate which was
originally two plates which have been fused together.” (Myriopods,
1895, p. 71.) That the segments were primitively separate is shown,
he adds, by the double nature of the circulatory system, the nerve
cord, and the first traces of segmentation in the mesoblast. Kenyon
believes that from the conditions in pauropods, Lithobius, etc.,
there are indications of alternate plates (not segments) having
disappeared, and of the remaining plates overgrowing the segments
behind them, so as to give rise to the anomalous double segments.[3]

[Illustration:

FIG. 11.—Sixth pair of legs of _Polyzonium germanicum_, ♀: _cs_,
ventral sacs; _cox_, coxa; _st_, sternal plate; _sp_,
spiracle.—After Haase.
]

Diplopods are also provided with eversible coxal sacs, in position like
those of Symphyla and Synaptera; Meinert, Latzel, and also Haase having
detected them in several species of Chordeumidæ, Lysiopetalidæ, and
Polyzonidæ (Fig. 11). In _Lysiopetalum anceps_ these blood-gills occur
in both sexes between the coxæ of the third to sixteenth pair of limbs.
In the Diplopods the blood-gills appear to be more or less permanently
everted, while in Scolopendrella they are usually retracted within the
body (Fig. 15, _cg_).

Diplopods also differ externally from insects in the genital armature, a
complicated apparatus of male claspers and hooks apparently arising from
the sternum of the sixth segment and being the modified seventh pair of
legs. In myriopods there are no pleural pieces or “pleurites,” so
characteristic of winged insects.

Perhaps the most fundamental difference between diplopods and insects is
the fact that the paired genital openings of the former are situated not
far behind the head between the second and third pair of legs. Both the
oviducts and male ejaculatory ducts are paired, with separate openings.
The genital glands lie beneath, while in chilopods they lie above the
intestine; this, as Korschelt and Heider state, being a more primitive
relation, since in Peripatus they also lie above the digestive canal.

The nervous system of diplopods is not only remarkable for the lack of
the tendency towards a fusion of the ganglia observable in insects, but
for the fact that the double segments are each provided with two
ganglia. The brain also is very small in proportion to the ventral cord,
the nervous system being in its general appearance somewhat as in
caterpillars.

The arrangement of the tracheæ and stigmata is much as in insects, but
in the Diplopoda the tracheary system is more primitive than in
chilopods, a pair of stigmata and a pair of tracheal bundles occurring
in each segment, while the bundles are not connected by anastomosing
branches, branched tracheæ only occurring in the Glomeridæ. The tracheæ
themselves are without spiral threads (tænidia). It is noteworthy that
the tracheæ arise much later than in insects, not appearing until the
animal is hatched; in this respect the myriopods approximate Peripatus.

In the Chilopoda also the parts of the head, except the epicranium, are
not homologous with those of insects, neither are the mouth-parts, of
which there are five pairs.

The structure of the head of centipedes is shown in part in Fig. 12,
compare also Fig. 8. It will be seen that it differs much from that of
the diplopods, though the mandibles (protomalæ) are homologous; they are
divided into a cardo and stipes, thus being at least two-jointed.

The second pair of postoral appendages is in centipedes very different
from the gnathochilarium of diplopods. As seen in Fig. 12 2, they are
separate, cylindrical, fleshy, five-jointed appendages, the maxillary
appendages of Newport, which are “connected transversely at their base
with a pair of soft appendages” (_c_), the lingua of Newport. The third
and fourth pair are foot-jaws, and we have called them _malipedes_, as
they have of course no homology with the maxillipedes of Crustacea. The
second pair of these malipedes, forming the last pair of
mouth-appendages, is the poison-fangs (4), which are intermediate
between the malipedes and the feet; Meinert does not allow that these
are mouth-appendages.

[Illustration:

FIG. 12.—Structure of a chilopod. _A_, _Lithobius americanus_, natural
size. _B_, under side of head and first two body-segments and legs,
enlarged: _ant_, antenna; 1, jaws; 2, first accessory jaw; _c_,
lingua; 3, second accessory jaw and palpus; 4, poison-jaw. (Kingsley
del.) _C_, side view of head (after Newport): _ep_, epicranium; _l_,
frontal plate; _sc_, scute; 1, first leg; _sp_, spiracle.
]

The embryology of Geophilus by Metschnikoff shows plainly the four pairs
of post-antennal appendages. The embryo Geophilus is hatched in the form
of the adult, having, unlike the diplopods, no metamorphosis, its
embryological history being condensed or abbreviated. But in examining
Metschnikoff’s figures certain primitive diplopod features are revealed.
The body of the embryo shortly before hatching is cylindrical; the
sternal region is much narrower than in the adult, hence the insertions
of the feet are nearer together, while _the first six pairs of
appendages begin to grow out before the hinder ones_. Thus the first six
pairs of appendages of the embryo Geophilus correspond to the antennæ,
two pairs of jaws, and three pairs of legs of the larval Julus. These
features appear to indicate that the chilopods may be an offshoot from
the diplopod stem. The acquisition of a second pair of legs to a segment
in diplopods, as in the phyllopod Crustacea, is clearly enough a
secondary character, as shown by the figures of Newport in his memoir on
the development of the Myriopoda (Pl. IV.). Thus the tendency in the
Myriopoda, both diplopods and chilopods, is towards the multiplication
of segments and the elongation of the body, while in insects the
polypodous embryo has the three terminal segments of the abdomen well
formed, these being, however, before hatching, partly atrophied, so that
the body of insects after birth tends to become shortened or condensed.
This indicates the descent of insects from ancestors with elongated
polypodous hind-bodies like Scolopendrella. Korschelt and Heider suggest
that the stem-form of myriopods was a homonomously jointed form like
Peripatus, consisting of a rather large number of segments, but we
might, with Haase, consider that the great number of segments which we
now find indicates a late acquisition of this form.

The genital opening in chilopods is single, and situated in the
penultimate segment of the body, as in insects. While recognizing the
close relationship of the Myriopoda with the insects, it still seems
advisable not to unite them into a single group (as Oudemans, Lang, and
others would do), but to regard them as forming an equivalent class. On
the other hand, when we take into account the form and structure of the
head, antennæ, and especially the shape of the first pair of
mouth-appendages, being at least two-jointed in both groups, we think
these characters, with the homonomously segmented body behind the head,
outweigh the difference in the position of the genital outlet, important
as that may seem. It should also be taken into account that while
insects are derived from polypodous ancestors, no one supposes, with the
exception of one or two authors, that these ancestors are the Myriopoda,
the latter having evidently descended from a six-legged ancestor, quite
different from that of the Campodea ancestor of insects.[4]

In regard to the sexual openings of worms, though their position is in
general in the anterior part of the body, it is still very variable,
though, in general, paired. In the oligochete worms the genital zone,
with the external openings, is formed by the segments lying between the
9th and 14th rings, though in some the genital organs are situated still
nearer the head. The myriopods, which evolved from the worms earlier
than insects, appear to have in their most primitive forms (the
Diplopoda) retained this vermian position of the genital outlets. In the
later forms, the chilopods, the genital openings have been carried back
to near the end of the body, as in insects. From observations made by
three different observers on the freshly hatched larva of the Julidæ, it
appears that the ancestral diplopods were six-footed, or oligopod, the
larva of Pauropus (Fig. 13) approaching nearest to our idea of the
ancestral myriopod, which might provisionally be named Protopauropus.

=Relations of the Symphyla to Insects.=—Opinions respecting the position
of the Symphyla, represented by Scolopendrella (Fig. 14), are very
discordant. By most writers since Newport, Scolopendrella has been
placed among the myriopods. The first author, however, to examine its
internal anatomy was Menge (1851), who discovered among other structures
(tracheæ, etc.) the silk-glands situated in the last two segments, and
which open at the end of each cercus. He regarded the form as “the type
of a genus or family intermediate between the hexapod Lepismidæ and the
Scolopendridæ.”

[Illustration:

FIG. 13.—_Pauropus huxleyi_, much enlarged. _A_, enlarged view of
head, antennæ, and first pair of legs (original). _B_, young.—After
Lubbock. _C_, longitudinal section of _Pauropus huxleyi_, ♂: _a_,
brain; _b_, salivary gland; _k_, mid-intestine; _g_, rectum; _h_,
ventral nerve-cord; _c_, bud-like remnants of coxæ; _d_, penis; _e_,
vesicula seminalis; _f_, ductus glandularis; _i_^1, divisions of
testes.—After Kenyon.
]

In 1873[5] the writer referred to this form as follows: “It may be
regarded as a connecting link between the Thysanura and Myriopoda, and
shows the intimate relation of the myriopods and the hexapods, perhaps
not sufficiently appreciated by many zoölogists.”

In 1880 Ryder regarded it as “the last survival of the form from which
insects may be supposed to have descended,” and referred it to “the new
ordinal group Symphyla, in reference to the singular combination of
myriopodous, insectean, and thysanurous characters which it
presents.[6]”

[Illustration:

FIG. 14.—_Scolopendrella immaculata_, from above,—after Lang; also
from beneath, the genital opening on the 4th trunk-segment: _sac_,
eversible or coxal sac; _an_, anus; _c_, cereopod; _v_, vestigial
leg.—After Haase, from Peytoureau. _A_ _B_ _C_, head and buccal
appendages of _Scolopendrella immaculata_: _A_, head seen from
above; _cl_, clypeus. _B_, head from beneath; _l_, first pair of
legs; _mx_, 1st maxilla; _mx_^1, 2d maxilla; _t_, “labial plates” of
Latzel, labium of Muhr. _C_, 1st maxilla; _l_, lacinia; _g_, galea;
_p_, rudiment of the palpus.—After Latzel. _D_, end of the body:
_p__{11}, eleventh, _p__{12}, twelfth undeveloped pair of legs;
_p__{13}, modified, vestigial legs, bearing tactile organs (_so_);
_sg_, cercopod, with duct of spinning gland, _dg_; _cd_, eversible
or coxal gland; _h_{8}s_, coxal spur of the 11th pair of legs.—After
Latzel from Lang.
]

Wood-Mason considered it to be a myriopod, and “the descendant of a
group of myriopods from which the Campodeæ, Thysanura, and Collembola
may have sprung.” We are indebted to Grassi for the first extended work
on the morphology of Scolopendrella (1885). In 1886 he added to our
knowledge facts regarding the internal anatomy, and gives a detailed
comparison with the Thysanura, besides pointing out the resemblances of
Scolopendrella to Pauropus, diplopods, chilopods, as well as Peripatus.

[Illustration:

FIG. 15.—Section of _Scolopendrella immaculata_: _œ_, œsophagus; _oe.
v_, œsophageal valve entering the mid-intestine (“stomach”); _i_,
intestine; _r_, rectum; _br_, brain; _ns_, abdominal chain of
ganglia; _ovd_, oviduct; _ov_, ovary; _s. gl_, silk-gland, and _op_,
its outer opening in cercus, _ur. t_, urinary tube; _cg_, coxal
glands or blood-gills.—Author _del._
]

In 1888 Grassi expressed his view as to the position of the Symphyla,
stating that it should not be included in the Thysanura, since it
evidently has myriopod characters; these being the supraspinal vessel,
the ventral position of the genital glands; the situation of the genital
opening in the fourth segment of the trunk, its ganglionic chain being
like that of diplopods, its having limbs on all the segments, etc. On
the other hand, Grassi has with much detail indicated the points of
resemblance to the Thysanura. The principal ones are the thin
integument, the want of sympathetic ganglia, the presence of a pair of
cephalic stigmata, like that said to occur in certain Collembola, and in
the embryo of Apis; two endoskeletal processes situated near the ventral
fascia of the head; the epicranial suture also occurring in Thysanura,
Collembola, Orthoptera, and other winged insects, and being absent in
diplopods and chilopods. He also adds that the digestive canal both in
Symphyla and Thysanura is divided into three portions; the malpighian
tubes in Thysanura present very different conditions (there being none
in Japyx), among which may be comprised those of Scolopendrella. In both
groups there is a single pair of salivary glands. The cellular
epithelium of the mid-intestine of Scolopendrella is of a single form as
in Campodea and Japyx. The fat-body, dorsal vessel, with its valves and
ostia, are alike in the two groups, as are the appendages of the end of
the abdomen, the anal cerci (cercopoda) of Scolopendrella being the
homologues of the multiarticulate appendages of Lepisma, etc., and of
the forceps of Japyx. In those of Scolopendrella, we have found the
large duct leading from the voluminous silk-gland, a single large sac
extending forwards into the third segment from the end of the body (Fig.
15, _s. gl_). Other points of resemblance, all of which he enumerates,
are the slight differences in the number of trunk-segments, the presence
in the two groups of the abdominal “false-legs” (parapodia), the dorsal
plate, and the mouth-parts. As regards the latter, Grassi affirms that
there is a perfect parallelism between those of Scolopendrella and
Thysanura. To this point we will return again in treating more
especially of those of the Symphyla. Finally, Grassi concludes that
there is “a great resemblance between the Thysanura and Scolopendrella.”
He, however, believed that the Symphyla are the forerunners of the
myriopods, and not of the insects, his genealogical tree representing
the symphylan and thysanuran phyla as originating from the same point,
this point also being, rather strangely, the point of origin of the
arachnidan phylum.

Haase (1889) regarded Scolopendrella as a myriopod, and Pocock (1893)
assigned the Symphyla to an independent class, regarding Scolopendrella
as “the living form that comes nearest to the hypothetical ancestor of
the two great divisions of tracheates.” Schmidt’s work (1895) on the
morphology of this genus is more extended and richly illustrated than
Grassi’s, his method of research being more modern. He also regards this
form as one of the lower myriopods.

In conclusion, it appears to us that, on the whole, if we throw out the
single characteristic of the anteriorly situated genital opening, the
ovarian tubes being directed toward the end of the body (Fig. 15, _ovd_,
_ov_), there is not sufficient reason for placing the Symphyla among the
Myriopoda, either below or near the diplopods. This is the only valid
reason for not regarding Scolopendrella as the representative of a group
from which the insects have descended, and which partly fills the wide
abyss between Peripatus and insects. With the view of Pocock, that both
insects and myriopods have descended from Scolopendrella, we do not
agree, because this form has so many insectean features, and a single
unpaired genital opening. For the same reason we should not agree with
Schmidt in interpolating the Symphyla between the Pauropoda and
Diplopoda. In these last two progoneate groups the genital openings are
paired, hence they are much more primitive types than Scolopendrella, in
which there is but a single opening. It seems most probable that the
Symphyla, though progoneate, are more recent forms than the progoneate
myriopods, which have retained the primitive feature of double sexual
outlets. It is more probable that the Symphyla were the descendants of
these polypodous forms. Certainly Scolopendrella is the only extant
arthropod which, with the sole exception of the anteriorly situated
genital opening, fulfils the conditions required of an ancestor of
Thysanura, and through them of the winged insects. No one has been so
bold as to suggest the derivation of insects from either diplopods or
chilopods, while their origin from a form similar to Scolopendrella
seems not improbable. Yet Uzel has very recently discovered that
Campodea develops in some respects like Geophilus, the primitive band
sinking in its middle into the yolk, with other features as in
chilopods.[7] The retention of a double sexual opening in the diplopods
is paralleled by the case of Limulus with its double or paired sexual
outlets, opening in a pair of papillæ, as compared with what are
regarded as the generalized or more primitive Crustacea, which have an
unpaired sexual opening.

The following summary of the structural features of the Symphyla, as
represented by Scolopendrella, is based mainly on the works of Grassi,
Haase, and Schmidt, with observations of my own.

=Diagnostic or essential characters of Symphyla.=—_Head shaped as in
Thysanura (Cinura), with the Y-shaped tergal suture, which occurs
commonly in insects (Thysanura, Collembola, Dermaptera, Orthoptera,
Platyptera, Neuroptera, etc.), but is wanting in Myriopoda (Diplopoda
and Chilopoda); antennæ[8] unlike those of Myriopoda in being very long,
slender, and moniliform. Clypeus distinct. Labrum emarginate, with six
converging teeth. Mandibles 2–jointed, consisting of a vestigial stipes
and distal or molar joint, the latter with eight teeth. First maxillæ
with an outer and inner mala situated on a well-developed stipes; with a
minute, 1–jointed palpus. Second pair of maxillæ: each forming two
oblong flat pieces, median sutures distinct, with no palpi; these pieces
are toothed in front, and appear to be homologous with the two median
pieces of the gnathochilarium of Diplopoda. Hypopharynx? Epipharynx?_

_Trunk with from fifteen to sixteen dorsal, more or less free subequal
scutes, the first the smallest. Pedigerous segments twelve; also twelve
pairs of 5–jointed legs, which are of nearly equal length, the first
pair 4–, the others 5–jointed, all ending in two claws, as in Synaptera
and winged insects. A pair of 1–jointed anal cerci homologous with those
of Thysanura and Orthoptera, into each of which opens a large abdominal
silk-gland. Abdominal segments with movable styles or “pseudopods”
(“Parapodia” of Latzel and of Schmidt), like those of Campodea and
Machilis, and situated on the base of the coxal joint in front of the
ventral sac. Within the body near the base of each abdominal style is an
eversible coxal sac or blood-gill (Fig. 15, cg). The single genital
opening is on the fourth trunk-segment in both sexes (Fig. 15, indicated
by the arrow). The malpighian tubes (ur. t) are two in number, opening
into the digestive canal at the anterior end of the hind intestine; they
extend in front to the third or second segment from the head. They are
broad and straight at their origin, becoming towards the end very
slender and convoluted._

_The three divisions of the digestive tract are as in insects, the
epithelium of the mid-gut being histologically as in Campodea and Japyx;
rectal glands are present. A pair of very large salivary glands are
situated in the first to the fourth trunk-segments, consisting of a
glandular portion with its duct, which unite into a common duct opening
on the under side of the head, probably in the labium._

_But a single pair of stigmata is present, and these are situated in the
front of the head, beneath the insertion of the antennæ and within the
stipes of the mandibles; the tracheæ are very fine, without spiral
threads (tænidia), and mostly contained within the head, two fine
branches extending on each side into the second trunk-segment._

_After birth the body increases in length by the addition of new
segments at the growing point._

In respect to the nervous system, there are no diagnostic characters;
there are, however, not as many as two pairs of ganglia to a segment.
The brain is well developed, sending a pair of slender nerves to the
small eyes. The ganglia of the segment bearing the first pair of legs is
fused with the subœsophageal ganglion. Grassi was unable to detect a
true sympathetic system, but he suspects the existence of a very small
frontal ganglion.

The slender dorsal vessel, provided with ostia and valvules, pulsates
along the entire length of the trunk; an aorta passes into the head.

The internal genital organs of both sexes are paired, and extend along
the greater part of the trunk; in either sex they may be compared to two
long, slender, straight cords extending from the fourth to the tenth
pair of legs. The two oviducts do not unite before reaching the sexual
opening (Fig. 15, _ovd_).

The male sexual organs are more complicated than the feminine. The
paired testicular tubes lie in trunk-segments 6 to 12, on each side, and
partly under the intestinal canal, communicating with each other by a
cross-anastomosis situated under the intestine, and which, like the
testes, is filled with sperm. Of the paired seminal ducts (vas deferens)
in trunk-segment 4, each unites again into a thick tube, sending a blind
tube forward into the third segment. Under the place of union of the two
vasa deferentia arise the paired ductus ejaculatorii, which open beneath
in the uterus masculinus. The anterior blind ends of the vasa deferentia
form a sort of small paired vesiculæ seminales in which a great quantity
of ripe sperm is stored. The uterus masculinus is in its structure
homologous with the evaginable penis of Pauropus, Polyxenus, and some
diplopods, and the sexual opening has without doubt become secondarily
unpaired. The sexual opening is rather long and is closed by two
longitudinal folds. “In several respects the male sexual organs of
Scolopendrella are like those of Pauropus; in the last-named form we
have indeed an unpaired testis, but also in Scolopendrella we see the
beginning of such a singleness; namely, the presence of an anastomosis
uniting the two tubes, their communication by means of a transverse
connecting canal and a glandular structure in the epithelium forming
them. The male sexual organs of Pauropus differ only through a still
greater complication.” (Schmidt.)

Scolopendrella in habits resembles chilopods, being found in company
with Geophilus burrowing deep in light sand under leaves, or living at
the surface of the ground under sticks or stones. It is very agile in
its movements, and is probably carnivorous. It was considered by Haase
to be eyeless, but the presence of two ocelli has been demonstrated both
by Grassi and by Schmidt. Whether the pigment and corneous facet are
present is not certain. The embryology is entirely unknown (although
Henshaw reports finding a hexapodous young one), and it need not be said
that a knowledge of it is a very great desideratum. It is most probable
that the young is hexapodous, since the first pair of limbs are
4–jointed, all the rest 5–jointed; while Newport, and also Ryder,
observed specimens with nine, ten, eleven, and twelve pairs, and
Wood-Mason confirms their observations, “which prove that a pair of legs
is added at each moult,” and he concludes that the addition of new
segments “therefore takes place in this animal by the intercalation of
two at each moult between the antepenultimate and penultimate sterna, as
in the Chilognatha, and as also in some of the Chilopoda.”

There is but one family, Scolopendrellidæ, and a single genus,
Scolopendrella, which seems to be, like other archaic types,
cosmopolitan in its distribution.

Our commonest species is _S. immaculata_ Newport, which occurs from
Massachusetts to Cordova, Mexico, and in Europe from England to the
Mediterranean and Russia; Mr. O. F. Cook tells me he has found a
species in Liberia, West Africa. The other species are _S.
notacantha_ Gervais, Europe and Eastern United States; _S. nivea_
Scopoli (_S. gratiæ_ Ryder), Europe and United States; _S. latipes_
Scudder, Massachusetts.

LITERATURE ON SCOLOPENDRELLA

=Newport, George.= Monograph of the class Myriopoda, order Chilopoda.
(Trans. Linn. Soc. xix, pp. 349–439, 1 Pl., 1845.)

=Menge, A.= Myriapoden der Umgegend von Danzig. (Neuste Schriften der
naturforsch. Gesell. Danzig. iv, 1851.)

=Ryder, John H.= Scolopendrella as the type of a new order of
articulates (Symphyla). (Amer. Nat., May, 1880, xiv, pp. 375, 376.)

—— The structure, affinities, and species of Scolopendrella. (Proc.
Acad. Nat. Soc. Phil., pp. 79–86, 1881, 2 Figs.)

=Packard, A. S.= Scolopendrella and its position in nature. (Amer.
Nat., 1881, pp. 698–704, Fig.)

=Muhr, Jos.= Die Mundtheile von Scolopendrella und Polyzonium. Prag,
1882, 1 Pl.

=Mason, J. Wood.= Morphological notes bearing on the origin of
insects. (Trans. Ent. Soc. London, 1879, pp. 145–167, Figs.)

—— Notes on the structure, post-embryonic development, and systematic
position of Scolopendrella. (Ann. and Mag. Nat. Hist., July, 1883,
pp. 53–63.)

=Latzel, Robert.= Die Myriapoden der osterreichisch-ungarischen
Monarchie, ii, Wien, 1884, pp. 1–39, Pls.

=Haase, Erich.= Die Abdominalanhänge der Insekten mit Berücksichtigung
der Myriapoden. (Morph. Jahrbuch, xv, pp. 331–435, 2 Pls., 1889.)

=Schmidt, Peter.= Beiträge zur Kenntnis der niederen Myriapoden.
(Zeits. f. wissen. Zool., lix, pp. 436–510, 2 Pls., 1895.)

INSECTA (HEXAPODA)

We are now prepared to discuss the fundamental or essential characters
of the insects, including the wingless subclass (Synaptera), and the
winged (Pterygota).

=Diagnostic characters of insects.=—_Body consisting of not more than
twenty-one segments, which are usually heteronomous or of unequal size
and shape, arranged in three usually well-defined regions_; i.e. _a
head, thorax, and hind-body or abdomen. Head small and flattened or
rounded, composed of not less than six segments, and bearing, besides
the eyes, at least four pairs of jointed appendages_; i.e. _one pair of
antennæ, and three pairs of masticatory appendages, the distal or molar
portion of which is primarily divided into three divisions, supported on
a stipes and cardo, and in certain orders modified into piercing or
sucking structures. The head is composed of an epicranium, bearing a
distinct clypeus and labrum, with the epipharynx. Mandibles 1–jointed,
without a palpus and very generally with no, or uncertain, traces of a
lacinia and a stipes. Two pairs of maxillæ; the first pair separate,
usually 3–lobed, comprising a lacinia, galea, and palpifer, with a
palpus which is never more than 6–jointed. The second pair united to
form the labium or under lip, composed of two laciniæ fused together; in
the generalized forms with a rudimentary galea; bearing a pair of palpi,
never more than 4–jointed; with paraglossæ sometimes present._

(_A third pair of mouth-appendages situated between the antennæ and
mandibles in the embryo of Anurida, and Apis, and adult Campodea._)

_The epipharynx forming the roof of the mouth, and bearing gustatory
organs. Hypopharynx usually well developed, lying on the under side of
the mouth, just above the labium, and receiving the end of the salivary
duct._

_Eyes of two kinds: a pair of compound, and from two to three simple
eyes (ocelli)._

_The thorax consisting of three segments, the two latter segments in the
winged orders highly differentiated into numerous tergal and lateral
pieces and a single sternum; in the Synaptera the segments are
undivided. (In the higher Hymenoptera the basal abdominal segment
coalesced with the thorax.) Three pairs of legs, each foot ending in a
pair of claws. Two pairs of wings (except in the Synaptera), a pair to
each of the two hinder thoracic segments; the wings occasionally reduced
or wanting in certain adaptive forms, which, however, had winged
ancestors._

_Abdomen consisting at the most of from ten to twelve segments. No
functional abdominal legs except in the Thysanura, and in the larvæ of
Lepidoptera. A pair of 1– or many-jointed cercopods on the tenth
segment; and in certain forms a pair of styles on the ninth segment. In
certain orders an ovipositor or sting formed of three pairs of styliform
processes; in Collembola a single pair of processes forming the elater._

_The genital openings opisthogoneate, usually single, but paired in
Thysanura (Lepisma), Dermaptera, and Plectoptera (Ephemeridæ)._

_The digestive canal in the winged orders is highly differentiated, the
fore-intestine being divided into an œsophagus and proventriculus, the
hind-intestine into an ileum, colon, and rectum, with rectal glands._

_The nervous system consists of a well-developed brain, in the more
specialized orders highly complicated; no more than thirteen pairs of
ganglia, which may be more or less fused in the more specialized orders.
Three frontal ganglia, and a well-developed, sympathetic system
present._

_Stigmata confined (except possibly in Sminthurus) to the thorax and
abdomen, not more than ten pairs in all, and usually but nine pairs.
Tracheal system as a rule highly differentiated; invariably with
tænidia._

_Dorsal vessel with ostia and valvules; no arteries except the cephalic
aorta; no veins. After birth there is in the more specialized pterygote
orders a reduction in the number of terminal segments of the abdomen._

_Development either direct (Synaptera), or with an incomplete (with
nymph and winged or imaginal stages), or complete metamorphosis; in the
latter case with a larval, pupal, and imago stage._

The insects may be divided into two sub-classes,—the _Synaptera_, and
the winged orders, _Pterygota_, of Gegenbaur (1877), since the
differences between the two groups appear on the whole to be of more
than ordinal rank.

1. EXTERNAL ANATOMY

_a._ The regions of the body

The insects differ from other arthropods in that the body is divided
into three distinct regions,—the head, thorax, and abdomen, the latter
regions in certain generalized forms not always very distinctly
differentiated. The body behind the head may also conveniently be called
the trunk, and the segments composing it the trunk-segments.

In insects the head is larger in proportion to the trunk than in other
classes, notably the Crustacea; the thorax is usually slightly or
somewhat larger than the head, while the hind-body or abdomen is much
the larger region, as it consists of ten to eleven, and perhaps in the
Dermaptera and Orthoptera twelve, segments, and contains the mid- and
hind-intestine, as well as the reproductive organs.

When we compare the body of an insect with that of a worm, in which the
rings are distinctly developed, we see that in insects ring distinctions
have given way to regional distinctions. The segments lose their
individuality. It is comparatively easy to trace the segments in the
hind-body of an insect, as in this region they are least modified; so
with the thorax; but in the head of the adult insect it is impossible to
discover the primitive segments, as they are fused together into a sort
of capsule, and have almost entirely lost their individuality.

In general it may be said that the head contains or bears the organs of
sense and of prehension and mastication of the food; the thorax the
organs of locomotion; and the abdomen those of reproduction.

When we compare the body of a wasp or bee with that of a worm, we see
that there is a decided transfer of parts headward; this process of
cephalization so marked in the Crustacea likewise obtains in insects.
Also the two hinder regions of the body are, in a much greater degree
than in worms, governed by the brain, the principal seat of the
intelligence, which, so to speak, dominates and unifies the functions of
the body, both digestive, locomotive, and reproductive, as also those of
the muscles moving the different segments and regions of the body. To a
large extent arthropodan morphology and class distinctions are based on
the regional arrangement of the somites themselves. Thus in the process
of grouping of the segments into the three regions, some increase in
size, while others undergo a greater or less degree of reduction; one
segment being developed at the expense of one or more adjoining ones.
This principle was first pointed out by Audouin, and is called Audouin’s
law. It is owing to the greater development of certain segments and the
reduction of others, both of the body-segments and of the segments of
the limbs, that we have the wonderful diversity of form in the species
and genera, and higher groups of insects, as well as those of other
arthropods.

_b._ The integument (exoskeleton)

The skin or integument of insects consists, primarily, as in worms and
all arthropods, of an epithelial layer of cells called the _hypodermis_.
This layer secretes the cuticle, which is of varying thickness and
flexibility, and is usually very dense, impermeable, and light, compared
with the crust of the Crustacea, where the cuticle becomes heavy and
solid by the deposition of the carbonate and phosphate of lime. This is
due to the presence of a substance called by Odier chitin.[9] The
cuticle is thin, delicate, and flexible between the joints; it is
likewise so in such diaphanous aquatic larvæ as that of Corethra, and in
the gills of aquatic insects, also in the walls of the tracheæ and of
the salivary ducts. The cuticle thus forms a more or less solid crust
which is broken into joints and pieces (sclerites), forming supports for
the attachments of the muscles and serving to protect the soft parts
within.

=Chitin.=—If we allow an insect to soak for a long time in acids, or
boil it in liquid potassa or caustic potash, the integument is not
affected. The muscles and the other soft parts are dissolved, leaving
the cuticle clear and transparent. This insolubility of the cuticle is
due to the presence of chitin, the insoluble residue left after such
treatment. It also resists boiling in acids, in any alkalies, alcohol or
ether. The chemical formula is C_{15}H_{26}N_{2}O_{10}.[10]

“Chitin forms less than one-half by weight of the integument, but it
is so coherent and uniformly distributed that when isolated by
chemical reagents, and even when cautiously calcined, it retains its
original organized form. The color which it frequently exhibits is
not due to any essential ingredient; it may be diminished or even
destroyed by various bleaching processes.” (Miall and Denny.)

“The chemical stability of chitin is so remarkable that we might
expect it to accumulate like the inorganic constituents of animal
skeletons, and form permanent deposits. Schlossberger (Ann. d. chem.
u. pharm., bd. 98) has, however, shown that it changes slowly under
the action of water. Chitin kept for a year under water partially
dissolved, turned into a slimy mass, and gave off a peculiar smell.
This looks as if it were liable to putrefaction. The minute
proportion of nitrogen in its composition may explain the complete
disappearance of chitin in nature.” (Miall and Denny, The Cockroach,
p. 29.)

Chitin, or a substance closely similar to it, occurs in worms and in
their tubes, especially in the pharyngeal teeth of annelids and in
their setæ. The shell of Lingula and the pen of cuttle-fish contain
true chitin (Krukenberg). The integument of Limulus, of trilobites,
and of Arachnida, as well as Myriopoda, appears to consist of
chitin.[11]

The chitin is rapidly deposited at the end of embryonic life, also
during the larval and pupal stages. As is well known, insects after
moulting are white, but in a few hours turn dark, and those which live
in total darkness are white, showing that light has a direct effect in
causing the dark color of the integument.

Moseley analyzed one pound weight of Blatta, and found plenty of iron
with a remarkable quantity of manganese.

Schneider regarded chitin as a hardening of the protoplasm rather than a
secretion, and the cuticle is looked upon as an exudation. It is
structureless, not consisting of cells, and consists of fine irregular
laminæ. “A cross-section of the chitinous layer or ‘cuticle’ examined
with a high power shows extremely close and fine lines perpendicular to
the laminæ.” In the cockroach the free surface of the cuticle is divided
into polygonal, raised spaces or areas which correspond each to a
chitinous cell of the hypodermis. (Miall and Denny.)

Numerous pore-canals pass through the cuticle of all the external parts
of the body. The larger canals nearly always form the way for the
passage of secretions from dermal cells, or connect with the cavities of
hairs or setæ; when very fine and not connected with hairs or scales,
they are either empty or filled with air, and may possibly serve for
respiration.

Vosseler distinguishes in the cuticle two layers of different physical
and chemical characters. Besides the external chitinous layer there is
an inner layer which entirely agrees with cellulose. (Zool.
Centralblatt, ii, 1895, p. 117.)

The reparative nature of chitin is seen in the fact that Verhoeff finds
that a wound on an adult Carabus, and presumably on other insects, is
speedily closed, not merely by a clot of blood, but by a new growth of
chitin.

_c._ Mechanical origin and structure of the segments (somites,
arthromeres, metameres, zonites)

The segments are merely thickenings of the skin connected by folds or
duplications of the integument, and not actually separate or individual
rings or segments. This is shown by longitudinal (sagittal) sections
through the body, and also by soaking or boiling the entire insect in
caustic potash, when it is seen that the integument is continuous and
not actually subdivided into separate somites or arthromeres, since they
are seen to be connected by a thin intersegmental membrane (Fig. 16).
But this segmentation or metamerism of the integument is, however, the
external indication of the segmentation of the arthropodan body most
probably inherited from the worms, being a disposition of the soft parts
which is characteristic of the vermian type. This segmentation of the
integument is correlated with the serial repetition of the ganglia of
the nervous system, of the ostia of the dorsal vessel, the primitive
disposition of the segmental and reproductive organs, of the soft,
muscular dissepiments which correspond to the suture between the
segments, and with the metameric arrangement of the muscles controlling
the movements of the segments on each other, and which internal
segmentation or metamerism is indicated very early in embryonic life by
the mesoblastic somites.

[Illustration:

FIG. 16.—Diagram of the anterior part of an insect, showing the
membranous intersegmental folds, _g_.—After Graber.
]

In the unjointed worms, as Graber states, the body forms a single but
flexible lever. In the earthworm the muscular tube or body-wall is
enclosed by a stiffer cuticle, divided into segments; hence the worm can
move in all required directions, but only by sections, as seen in Fig.
16, which represents the thickened integument divided into segments, and
folded inward between each segment, this thin portion of the skin being
the intersegmental fold. Each segment corresponds to a special zone of
the subdivided muscular tube (_m_), the fascia extending longitudinally.
The figure shows the mode of attachment of the fascia of the muscle-tube
to the segment. The anterior edge is inserted on the stiff, unyielding,
inner surface of each segment: the hinder edge of the muscle is attached
to the thin, flexible, intersegmental fold, which thus acts as a tendon
on which the muscle can exert its force. (Graber.)

[Illustration:

FIG. 17.—Diagram of the integument and arrangement of the segmental
muscles: _A_, relaxed; _m_, muscle; _g_, membranous articulation;
_r_, chitinous ring. _B_, the same contracted on both sides. _C_, on
one side.—After Graber.
]

“Fig. 17 makes this still clearer. The muscles (_m_) extend between two
segments immediately succeeding each other. Supposing the anterior one
(_A_) to be stationary, what do we then see when the muscle contracts?
Does it also become shorter? The intersegmental fold is drawn forwards,
and hence the entire hinder segment moves forward and is shoved into the
front one, and so on with the others, as at _B_. Afterwards, if the
strain of the muscle is relieved by the diminishing action of the
tensely stretched, intersegmental membrane, it again returns to a state
of rest.” (Graber.)

[Illustration:

FIG. 18.—Diagrams to demonstrate the mechanism of the motion of the
segmented body in the Arthropoda: One larger segment (_cf_) and 4
smaller. The exoskeleton is indicated by black lines, the
interarticular membranes by dotted lines. The hinges between
consecutive segments are marked _at_, tergal (dorsal) skeleton; _s_,
sternal (ventral) skeleton; _d_, dorsal longitudinal muscles =
extensors (and flexors in an upward direction); _v_, ventral
longitudinal muscles = flexors. In _B_, the row of segments is
stretched; in _A_, by the contraction of the muscles (_d_) bent
upward; in _C_, downward; _tg_, tergal; _sg_, sternal interarticular
membranes.—After Lang.
]

While we look upon the dermal tube of worms as a single but flexible
lever, the body of the arthropods, as Graber states, is a linear system
of stiff levers. We have here a series of stiff, solid rings, or hooks,
united by the intersegmental membrane into a whole. When the muscles,
extending from one ring to the next behind contract, and so on through
the entire series, the rings approximate each other.

The ectoskeletal segments bend to one side by the contraction of the
muscles on one side, the point of the outer segmental fold opposite the
fixed point becoming converted into the turning-point (_C_).

The usual result of the arrangement of the locomotive system is the
simple curving of the body (_C_), and then the alternate bending of the
body to right and left, which produces the serpentine movements
characteristic of the earthworms, the centipede, and many insect larvæ.
The most striking example of the wonderful variety of movements which
can be made by an insect are those of the Syrphus larva. When feeding
amid a herd of aphides, it is seen to now raise the front part of the
body erect and stiff, then to bend it down, or rapidly turn it to either
side, or move it in a complete circle. (Graber, pp. 23–26.)

The arrangement and mode of working of the muscles, says Lang, is
illustrated by Fig. 18, which shows us five segments, one larger (_ct_)
and four smaller, in vertical projection. The thicker portion of the
integument is marked by strong outlines, the delicate and flexible
interarticular membranes (_tg_, _sg_) in dotted lines. The hinges
between two consecutive segments are marked _a_. A dorsal muscle (_d_)
is attached to the larger segment (_ct_), and runs through the smaller
segments, being inserted in the dorsal portion of the crust (_t_) of
each by means of a bundle of fibres. A ventral muscle (_v_) does the
same on the sternal side (_s_).

“The skeletal segments,” adds Lang, “may be compared to a double-armed
lever, whose fulcrum lies in the hinges. If the dorsal muscle contracts,
it draws the dorsal arm of the lever (the tergal portion of the
skeleton) in the direction of the pull towards the larger segments; the
tergal interarticular membranes become folded, the ventral stretched,
and the four segments bend upward (Fig. 18, _A_). If the ventral muscle
contracts, while at the same time the dorsal slackens, the row of
segments will be bent downwards (Fig. 18, _C_).”

L. B. Sharp suggests, that in the Crustacea the rings formed by “the
regularity and stress of muscular action” would be hardened by the
deposition of lime at the most prominent portion, _i.e._ between what we
have called the intersegmental folds. (American Naturalist, 1893, p.
89.) Cope also states that “with the beginning of induration of the
integument, segmentation would immediately appear, for the movements of
the body and limbs would interrupt the deposit at such points as would
experience the greatest flexure. The muscular system would initiate the
process, since flexure depends on its contractions, and its presence in
animals prior to the induration of the integuments in the order of
phylogeny, furnishes the conditions required.” (The Primary Factors of
Organic Evolution, p. 268, 1895.)

It is apparent that the jointed or metameric structure of the bodies of
insects and other arthropods is an inheritance from the segmented worms.
In the worms the body is a continuous dermo-muscular tube, while in
arthropods this tube is divided into regions, and the cuticle is thicker
and more resistant. To go back to the incipient stages in the process of
segmentation of the body, we conceive that the worms probably arose from
a creeping gastrula-like form, the gastræa. The act of creeping
gradually induced an elongated shape of the body. The movement of such
an organism in a forward direction would gradually evolve a fore and
aft, dorsal and ventral, and bilateral symmetry. As soon as this was
attained, as the effect of creeping over rough irregular surfaces there
would result mechanical lateral strains intermittently acting during the
serpentine movements of the worm. The integument would, we can readily
suppose, tend to bend or yield, or become permanently wrinkled, at more
or less regular intervals. The arrangement of the muscles would
gradually conform to this habit of creeping, and finally the nervous
system and other organs more directly connected with the creeping
movements of the organism would tend to be correlated in their
arrangement with that of the segments. In this way the homonomous
segments of the annelid body probably became developed, and their
relations and shapes were eventually fixed by inheritance. After this
stage was reached, and limbs began to appear, the segments would tend to
become heteronomous, and to be grouped into regions.

[Illustration:

FIG. 19.—_Dujardinia rotifera_, with jointed tentacles and caudal
appendages.—With some changes, after Quatrefages.
]

The origin of the joints or segments in the limbs of arthropods was
probably due to the mechanical strains to which what were at first soft
fleshy outgrowths along the sides of the body became subjected. Indeed,
certain annelid worms of the family Syllidæ have segmented tentacles and
parapodia, as in Dujardinia (Fig. 19). We do not know enough about the
habits of these worms to understand how this metamerism may have arisen,
but it is possibly due to the act of pushing or repeated efforts to
support the body while creeping over the bottom among broken shells,
over coarse gravel, or among seaweeds.

It is obvious, however, that the jointed structure of the limbs of
arthropods, if we are to attempt any explanation at all of the origin of
such structure, was primarily due mainly to lateral strains and impacts
resulting from the primitive endeavors of the ancestral arthropods to
raise and to support the body while thus raised, and then to push or
drag it forward by means of the soft, partially jointed, lateral limbs
which were armed with bristles, hooks, or finally claws.

On the other hand, by adaptation, or as the result of parasitism and
consequent lack of active motion, the original number of segments may by
disuse be diminished. Thus in adult wasps and bees, the last three or
four abdominal segments may be nearly lost, though the larval number is
ten. During metamorphosis the body is made over, and the number, shape,
and structure of the segments greatly modified. In the female of the
Stylopidæ the thorax loses all traces of segments, and is fused with the
head, and the abdominal segments are faintly marked, losing their
chitin.

While the maxillæ have several joints, the mandibles are 1–jointed, but
there are traces of two joints in Campodea, certain beetles, etc. In the
antenna there is a great elasticity in respect to the number of joints,
which vary from one or two to a hundred or more. It is likewise so in
the thoracic legs, where the number of tarsal joints varies from one to
five; also in the cercopoda, the number of joints varying from one or
two to twelve or more.

_d._ Mechanical origin of the limbs and of their jointed structure

We have already hinted at the mode of origin of the limbs of arthropods.
Like the body or trunk, the limbs are chitinous dermo-muscular tubes,
with a dense solid cuticle, and internal muscles, and were it not for
their division at more or less regular intervals into segments, forming
distinct sets of levers, set up by the strains in these tubular
supports, there would be no power of varied motion.

Even certain worms, as already stated, have their tentacles and
parapodia, or certain appendages of their parapodia, more or less
jointed, but there are no indications of claws or of any other hard
chitinous armature at the extremity, and the skin is thin and soft.

In the most simple though not the most primitive arthropods, such as the
Tardigrades, whose body is not segmented, there are four pairs of short
unjointed legs, ending each in two claws, which have probably arisen in
response to the stimulus of pushing or dragging efforts.

The legs of Peripatus are unjointed, and have a thin cuticle, but end in
a pair of claws, which have evidently arisen as a supporting armature,
the result of the act of moving or pulling the body over the uneven
surface of the ground.

[Illustration:

FIG. 20.—A prothoracic leg of Chironomus larva; and pupa.
]

[Illustration:

FIG. 21.—_A_, larva of _Ephydra californica_: _a_, _b_, _c_, pupa.
]

There is good reason to suppose that such limbs arose from dynamical
causes, similar to those exciting the formation of secondary adaptations
such as are to be seen in the prop or supporting legs of certain
dipterous larvæ, as the single pair of Chironomus (Fig. 20) and
Simulium, or the series of unjointed soft tubercles of Ephydra (Fig.
21), etc., which are armed with hooks and claws, and are thus adapted
for dragging the insect through or over vegetation or along the ground.

Now by frequent continuous use of such unjointed structures, the cuticle
would tend to become hard, owing to the deposit of a greater amount of
chitin between the folds of the skin, until finally the body being
elongated and homonomously segmented, the movements of walking or
running would be regular and even, and we would have homonomously
jointed legs like those of the trilobites, or of the most generalized
Crustacea and of Myriopoda.

In the most primitive arthropods,—and such we take it were on the whole
the trilobites, rather than the Crustacea,—the limbs were of nearly the
same shape, being long and slender and evenly jointed from and including
the antennæ, to the last pair of limbs of the abdominal region. In these
forms there appear to be, so far as we now know, no differentiation into
mandibles, maxillæ, maxillipedes, and thoracic legs, or into gonopoda.
The same lack of diversity of structure and function of the
head-appendages has survived, with little change, in Limulus. In the
trilobites (Fig. 1) none of the limbs have yet been found to end in
claws or forceps; being in this respect nearly as primitive as in the
worms. Secondary adaptations have arisen in Limulus, the cephalic
appendages being forcipated, adapted as supports to the body and for
pushing it onward through the sand or mud, while the abdominal legs are
broad and flat, adapted for swimming and bearing the broad gill-leaves.

It is thus quite evident that we have three stages in the evolution of
the arthropodan limb; _i.e._ 1, the syllid stage, of simple, jointed,
soft, yielding appendages not used as true supports (Fig. 19); 2, the
trilobite stage, where they are more solid, evenly jointed, but not
ending in claws; and by their comparatively great numbers (as in the
trilobite, Triarthrus) fully supporting the body on the bottom of the
sea. In Limulus they are much fewer in number, thicker, and acting as
firm supports, the cephalic limbs of use in creeping, and ending in
solid claws. 3, The third stage is the long slender swimming
head-appendages of the nauplius stage of Crustacea.

As regards the evolution of limbs of terrestrial arthropods, we have the
following stages: 1, the soft unjointed limbs of Tardigrades, ending in
two claws, and those of Peripatus, and the pseudo- or prop-legs of
certain dipterous larvæ; 2, finally the evolution of the long, solid,
jointed limbs of Pauropus and other primitive myriopods, the legs
forming solid, firm supports elevating the body, and enabling the insect
to drag itself over the ground or to walk or run. When the body is
elongated and many-segmented, the legs are necessarily numerous; but
when it is short, the legs become few in number, _i.e._ six, in the
hexapodous young of myriopods and in insects, or eight in Arachnida.
Whenever the legs are used for walking, _i.e._ to raise and support the
body, they end in a solid point or in a pair of forceps or claws. On the
other hand, as in phyllopods, where the legs are used mainly for
swimming, they are unarmed and are soft and membranous, or, as in the
limbs of the nauplius or zoëa stage of crustaceans, end in a simple soft
point, which often bears tactile setæ.

The tarsal joints are more numerous in order to give greater flexibility
to the limb in seizing and grasping objects, both to drag the body
forwards and to support it.

Unlike those of the Crustacea, the limbs of insects are not primitively
biramose, but single, the three-lobed first maxillæ, and secondarily
bilobed second maxillæ being the result of adaptation. Embryology on the
whole proves the truth of this assumption; the maxillæ of both pairs are
at first single buds, afterwards becoming lobed. All the appendages of
the body, including the ovipositor or sting, are modified limbs, as
shown by their embryological development.

It is noticeable that in the crab, where the body is raised by the limbs
above the bottom, it is much shorter and more cephalized than in the
shrimps. Also in the simply walking and running spiders, the hind-body
is shorter than in scorpions, while in the running and flying insects,
such as the Cicindelidæ, and in the swiftly flying flies and bees, there
is a tendency to a shortening of the body, especially of the abdomen.
The long body of the dragon-fly is an impediment to flight, but
compensated for by the action of the large wings.

The arthropodan limb is a compound leverage system. It is, says Graber,
a lateral outgrowth of the trunk, which repeats in miniature that of the
main trunk, its single series of joints or segments forming a jointed
dermo-muscular tube. Yet the lateral appendages of an insect differ from
the main trunk in two ways: (1) they taper to the end which bears the
two claws, and (2) their segments are in the living animal arranged not
in a straight line, but at different angles to each other. The basal
joint turning on the trunk acts as the first of a whole series of
levers. The second joint, however, is connected with the musculature of
the first or basal joint, and thus each succeeding joint is moved on the
one preceding. Each lever, from the first to the last, is both an active
and a passive instrument. (Graber.)

While, however, as Graber states, the limbs possess their own sets of
muscles and can move by the turning of the basal joint, the labor is
very much facilitated, as is readily seen, by the trunk, though the
latter has to a great extent delegated its locomotive function to the
appendages, which again divide its labor among the separate joints.

Graber then calls attention to the analogy of the mechanics of
locomotion of insects to those of vertebrates. An insect’s and a
vertebrate’s legs are constructed on the same general mechanical
principles, the limbs of each forming a series of levers.

[Illustration:

FIG. 22.—Diagram of the knee-joint of a vertebrate (_A_) and an
insect’s limb (_B_): _a_, upper; _b_, lower, shank, united at _A_ by
a capsular joint, at _B_ by a folding joint; _d_, extensor or
lifting muscle; _d_^1, flexor or lowering muscle of the lower joint.
The dotted line indicates in _A_ the contour of the leg.—After
Graber.
]

Fig. 22, _A_, represents diagrammatically the knee joint of a
vertebrate, and _B_ that of an insect; _a_, the femur or thigh, and _b_,
the tibia or shank. In the vertebrate the internally situated bones are
brought into close union and bend by means of a hinge-joint; so also in
the chitinous-skinned insect.

The stiff dermal tube of the insect acts as a lever by means of the thin
intersegmental membrane (_c_) pushed in or telescoped in to the thigh
joint, a special joint-capsule being superfluous. The muscles are in
general the same in both types; they form a circle. In both the shank is
extended by the contraction of the upper muscles (_d_) and is bent by
the contraction of the lower (_d^1_). The intersegmental membrane of the
insect’s limb is in a degree a two-armed lever, whose pivot (_f_) lies
in the middle. The internal invagination of the intersegmental fold
(_B_, _g-h_) affords the necessary support to the muscles acting like
the tendon in the vertebrate. (Graber.)

[Illustration:

FIG. 23.—Primitive band or germ of a Sphinx moth, with the segments
indicated, and their rudimentary appendages: _c_, upper lip; _at_,
antennæ; _md_, mandibles; _mx_, _mx′_, first and second maxillæ;
_l_, _l′_, _l″_, legs; _al_, abdominal legs.—After Kowalevsky.
]

Graber also calls attention to the fact that this insect limb differs in
one important respect from that of land vertebrates. The leverage system
in the last is divided at the end into five parallel divisions or
digits. In arthropods, on the contrary, all the joints succeed one
another in a linear series.

In insects, as well as in other arthropods, modifications of the limbs
usually take the form of a simple reduction in the number of segments.
Thus while the normal number of tarsal joints is five, we have trimerous
and dimerous Coleoptera, and in certain Scarabæidæ the anterior tarsi
are lost.

Savigny was the first, in 1816, in his great work, “Théorie des organes
de la bouche des Crustacés et des Insectes,” to demonstrate that not
only were the buccal appendages of biting insects homologous with those
of bugs, moths, flies, etc., but that they were homologous with the
thoracic legs, and that thus a unity of structure prevails throughout
the appendages of the body of all arthropods. Oken also observed that
“the maxillæ are only repeated feet.”

What was modestly put forth as a theory by the French morphologist has
been abundantly proved by the embryology of insects of different orders
to be a fact. As shown in Fig. 23 the antennæ and buccal appendages
arise as paired tubercles exactly as the thoracic legs. The abdominal
region also bears similar embryonic or temporary limbs, all of which in
those insects without an ovipositor disappear, except the cercopoda,
after birth.

LITERATURE ON THE EXTERNAL ANATOMY

General

=Swammerdam, Johann.= Biblia naturæ. (In Dutch, German, and English.)
1737–1738, fol., London, 1758, Pls.

=Réaumur, Réné Antoine Ferchault, de.= Mémoires pour servir à
l’histoire des insectes. i-iv, 4º, Paris, 1734–1742.

=Lyonet, Pieter.= Traité anatomique de la chenille, qui ronge le bois
de saule, etc. 4º, pp. xxii, 616. À la Haye, 1732. Tab. 18.

—— Recherches sur l’anatomie et les metamorphoses de differentes
espèces d’insectes. Ouvrage posthume, publié par M. W. de Haan. pp.
580, tab. 54, 1832.

=Latreille, Pierre André.= Des rapports généraux de l’organization
extérieure des animaux invertébres articulés, et comparaison des
Annelides avec les Myriapodes. (Mémoires du Mus. d’Hist. Nat., 1820,
vi, pp. 116–144.)

—— De quelques appendices particuliers du thorax de divers insectes.
(Mémoires du Mus. d’Hist. Nat., 1821, vii, pp. 1–21).

—— Observations nouvelles sur l’organization extérieure et générale
des animaux articulés et à pieds articulés, et application de ces
connoissances à la nomenclature des principales parties des mêmes
animaux. (Mèmoires du Mus. d’Hist. Nat., viii, 1822, pp. 169–202.)

=Cuvier, George Leopold Christian Dagobert.= Rapport sur les
recherches anatomiques sur le thorax des animaux articulés et celui
des insectes en particulier par M. V. Audouin. 4º, pp. 15, tab. 1,
Paris, 1823.

=Audouin, Jean Victor.= Recherches anatomiques sur le thorax des
animaux articulés et celui des insectes hexapodes en particulier.
(Annales des Sciences naturelles, i, pp. 97–135, 416–432, 1824.)

=Kirby, William, and William Spence.= Introduction to entomology.
i-iv, 1816–1828, London.

=Straus-Durckheim, Hercule.= Considérations générales sur l’anatomie
comparée des animaux articulés. Paris, 1828, atlas of 19 plates.

=MacLeay, William Sharp.= Explanation of the comparative anatomy of
the thorax in winged insects, with a review of the present state of
the nomenclature of its parts. (Zoöl. Journal, v, pp. 145–179, 1830,
2 Pls.)

=Burmeister, Hermann.= A manual of entomology. Trans. by W. E.
Shuckard, 8º, pp. 654, London, 1836, 32 Pl.

=Westwood, John Obadiah.= An introduction to the modern classification
of insects, i, ii, 8º, pp. 462, 587, 158, 1 Pl. and 133 blocks of
figs., 1839–1840.

=Newport, George.= Art. Insecta in Todd’s Cyclopædia of Anatomy and
Phys. ii, pp. 853–994, 1839, Figs. 329–439.

=Erichson, Wilhelm Ferdinand.= Entomographien. Berlin, 1840.

=Brullé, Auguste.= Recherches sur les transformations des appendices
dans les articulés. (Annales des Sciences nat. Sér. 3, ii, pp.
271–374, tab. 1, 1844.)

=Winslow, A. P.: son.= Om byggnaden af thorax hos Insekterna.
Helsingborg, 1862, 1 Pl, pp. 24.

=Packard, Alpheus Spring.= Guide to the study of insects. 1869.

—— Systematic position of the Orthoptera in relation to other insects.
(Third report U. S. Ent. Commission, pp. 286–345, 1883, Pls.
xxiii-lxi.)

=Graber, Vitus.= Die Insekten. 12º, pp. 403, 603, München, 1877, many
Figs.

=Huxley, Thomas Henry.= A manual of the anatomy of invertebrated
animals. 12º, pp. 397–451, Figs., London, 1877.

=Hammond, Arthur.= Thorax of the blow-fly. (Journ. Linn. Soc., London,
xv. Zoöl., 1880, pp. 31.)

=Brauer, Friedrich.= Ueber das Segment médiaire Latreille’s. (Sitzb.
d. k. Akad. d. Wissensch. Wien, 1882, pp. 218–241, 3 tab.)

—— Systematisch-zoologische Studien. (Ibid., 1885, pp. 237–413.)

=Gosch, C. C. A.= On Latreille’s theory of “Le Segment médiaire.”
(Nat. Tidsskrift (3), xiii, pp. 475–531, 1883.)

=Miall, L. C., and Denny, Alfred.= The structure and life-history of
the cockroach (_Periplaneta orientalis_). An introduction to the
study of insects. 8º, pp. 224, London, 1886.

=Cheshire, Frank R.= Bees and bee-keeping. i, Scientific, London,
1886, Pls. and Figs.

=Lang, Arnold.= Text-book of comparative anatomy. i, pp. 426–508,
1891, many Figs.

=Kolbe, H. J.= Einführung in die Kenntniss der Insekten. 8º, pp. 709,
324 figs., Berlin, 1893.

=Sharp, David.= The Cambridge natural history. Insecta, i, 8º, pp.
83–584, 1895, Figs. 47–371.

Also the works of Bos, Chabrier, Cholodkowsky, Comstock, Dewitz, Eaton,
Erichson, Gerstaecker, Girard, Grassi, Hagen, Haase, Kellogg, Knoch,
Lacordaire, Latreille, Leuckart, Lendenfeld, Lowne, Lubbock, Mayer,
Meinert, F. Müller, Osten-Sacken, Pagenstecher, Reinhard, Schaum,
Schiödte, Scudder, J. B. Smith, Spinola, Stein, Weismann, Wood-Mason.

THE HEAD AND ITS APPENDAGES

_a._ The head

[Illustration:

FIG. 24.—Presumed larva of Nemoptera (_Necrophilus arenarius_),
Pyramids of Egypt.—After Roux, from Sharp.
]

While the head is originally composed of probably not less than six
segments, these are in the adult insect fused together into a capsule or
hard chitinous box, the _epicranium_, with no distinct traces of the
primitive segments. The head contains the brain and accessory ganglia,
the mouth or buccal cavity, also the air-sacs in many winged forms, and
gives support to the external organs of sense, the antennæ, and to the
buccal appendages, the larger part of the interior being filled with the
muscles moving these structures. The solid walls of the head serve as a
lever or support for the attachment of these muscles, especially those
of the mandibles. Thus there is a correlation between the large size of
the mandibles of the soldier white ants and ants, the head being
correspondingly large to accommodate the great mandibular muscles. The
other extreme is seen in the larva of Necrophilus (Fig. 24), with its
long slender neck and diminutive head.

=The clypeus.=—This is that part of the head situated in front of the
epicranium, and anterior to the eyes, forming the roof of the posterior
part of the mouth, and is, as embryology shows, probably a tergal
sclerite. It varies greatly in shape and size in the different orders of
insects. It is often divided into two parts, the _clypeus posterior_ and
_clypeus anterior_, or which may be designated as the _post-_ and
_ante-clypeus_ (Figs. 29, _B_).

=The labrum.=—The “upper lip” or labrum is an unpaired flap-like piece
hinged to the front edge of the clypeus, and may be seen to move up and
down when the insect moves its mandibles. It forms the roof of the
anterior part of the mouth (Figs. 69, 74), and its inner side is lined
with a soft membrane, usually provided with hairs and sense-papillæ or
cups, forming the epipharynx.

The labrum is more or less deeply bilobed, especially in caterpillars
and in adult Staphylinidæ, and has been thought by some writers
(Kowalevsky, Carrière, and also Chatin) to represent a pair of
appendages, but Heymons (1895) refutes this view, stating as his reason
that the labrum arises between the two halves of the nervous system
(protocerebrum), while all the true appendages arise on each side of the
nervous system. (See also Fig. 34.)

[Illustration:

FIG. 25.—Front view of the head of _C. spretus_: _E_, epicranium; _C_,
clypeus; _L_, labrum; _O_, _o_, ocelli; _e_, eye; _a_, antenna;
_md_, mandible; _mx_, portion of maxilla uncovered by the labrum;
_p_, maxillary palpus; _p′_, labial palpus.
]

In the fleas (Siphonaptera) both the clypeus and labrum are wanting.

While it apparently forms an anterior specialized portion of the
procephalic lobes, Viallanes regarded it as belonging to the third, or
his tritocerebral, segment, since the labral nerves arise from the
tritocerebral ganglia. But since in all the early as well as late stages
of embryonic life it appears to be situated in front of the mouth, it
would seem to belong to the first segment.

In the embryo of Blatta it first appears as a thick crescentic fold
being slightly divided anterior to the mouth, and in Doryphora it
appears as a heart-shaped or deeply bilobed prominence situated in front
of the mouth (Wheeler).

=The epipharynx and labrum-epipharynx.=—The epipharynx is the under
surface or pharyngeal lining of the clypeus and labrum, forming the
membranous roof of the mouth. As it contains the organs of taste and has
been generally overlooked by entomologists, we may dwell at some length
on its structure in different orders.

Réaumur was, so far as we have been able to ascertain, the first author
to describe and figure the epipharynx, which he observed in the honey
bee and bumble bee, and called _la langue_, remarking that it closes the
opening into the œsophagus, and that it is applied against the palate.
According to Kirby and Spence, De Geer described the epipharynx of the
wasp; and Latreille referred to it, calling it the _sous labre_.

The name _epipharynx_ was bestowed upon this organ by Savigny, who thus
speaks of that of the bees: “Ce pharynx est, à la vérité, non seulement
caché par la lèvre supérieure, mais encore exactement recouvert par un
organe particulier que Réaumur a déjà décrit. C’est une sorte
d’appendice membraneux qui est reçu entre les deux branches des
mâchoires. Cette partie ayant pour base le bord supérieur du pharynx,
peut prendre le nom d’_épipharynx_ ou d’_épiglosse_.”

He also describes that of Diptera. What Walter has lately proved to be
the epipharynx of Lepidoptera was regarded by Savigny and all subsequent
writers as the labrum.

The latest account of the function of this organ is that by Cheshire,
who states that the tube made by the maxillæ and labial palpi cannot act
as a suction pipe, because it is open above. “This opening is closed by
the front extension of the epipharynx, which closes down to the maxillæ,
fitting exactly into the space they leave uncovered, and thus the tube
is completed from their termination to the œsophagus.”

[Illustration:

FIG. 26.—Epipharynx of _Phaneroptera angustifolia_: _cl_, clypeus;
_lbr. e_, labrum-epipharynx; _t_ _c_, taste cups, both on the
clypeal and on the labral regions.
]

[Illustration:

FIG. 27.—Epipharynx of _Hadenœcus subterraneus_, cave cricket.
]

It is singular that this organ is not mentioned in Burmeister’s Manual
of entomology, in Lacordaire’s Introduction à l’entomologie, or by
Newport in his admirable article _Insecta_ in Todd’s Cyclopedia of
anatomy. Neither has Straus-Durckheim referred to or figured it in his
great work on the anatomy of _Melolontha vulgaris_.

In their excellent work on the cockroach, Miall and Denny state that
“The epipharynx, which is a prominent part in Coleoptera and Diptera, is
not recognizable in Orthoptera” (p. 45). We have, however, found it to
be always present in this order (Figs. 26, 27).

We are not aware that any modern writers have described or referred to
the epipharynx of the mandibulate orders of insects. Although Dr. G.
Joseph speaks of finding taste-organs on the palate of almost every
order of insects, especially plant-feeding forms, we are unable to find
any specific references, his detailed observations being apparently
unpublished.

The epipharynx is so intimately associated with the elongated labium of
certain Diptera, that, with Dr. Dimmock, we may refer to the double
organ as the labrum-epipharynx; and where, as in the lepidopterous
_Micropteryx semipurpurella_, described and figured by Walter, and the
Panorpidæ (Panorpa and Boreus), the labrum seems pieced out with a thin,
pale membranous fold which appears to be an extension of the epipharynx,
building up the dorsal end of the labrum, this term is a convenient one
to use.

In the lower orders of truly mandibulate insects, from the Thysanura to
the Coleoptera, excluding those which suck in liquid food, such as the
Diptera, Lepidoptera, and Hymenoptera, and the Mecoptera (Panorpidæ)
with their elongated head and feeble, small mandibles, the epipharynx
forms a simple membranous palatal lining of the clypeus and labrum. In
such insects there is no soft projecting or pendant portion, fitted to
close the throat or to complete a partially tubular arrangement of the
first and second maxillæ.

In all the mandibulate insects, then, the epipharynx forms simply the
under surface or pharyngeal lining of the clypeus and labrum, the
surface being uniformly moderately convex, and corresponding in extent
to that of the clypeus and labrum, posteriorly merging into the palatal
wall of the pharynx; the armature of peculiar gathering-hairs sometimes
spreading over its base, being continuous with those lining the mouth
and beginning of the œsophagus. The suture separating the labrum from
the clypeus does not involve the epipharynx, though since certain
gustatory fields lie under the front edge of the clypeus, as well as
labrum, one may in describing them refer to certain fields or groups of
cups or pits as occupying a labral or clypeal region or position.

The lack of traces of a suture in the epipharynx corresponding to the
labral suture above, suggests that the labrum does not represent a pair
of coalesced appendages, and that it, with the clypeus, simply forms the
solid cuticular roof of the mouth.

The only soft structures seen between the epipharynx and labrum, besides
the nerves of special sense, are the elevator muscles of the labrum, and
two tracheæ, one on each side.

The structure and armature of the epipharyngeal surface even besides the
taste-pits, taste-cups and rods, is very varied, the setæ assuming very
different shapes. There seem to be two primary forms of setæ, (1) the
normal forms which arise from a definite cell; and (2) soft, flattened,
often hooked hairs which are cylindrical towards the end, but arise from
a broad triangular base, without any cell-wall. These are like the
“gathering hairs” of Cheshire, situated on the bees’ and wasps’ tongue;
they also line the walls of the pharynx and extend toward the œsophagus.
They are also similar to the “hooked hairs” of Will. The first kind, or
normal setæ, are either simply defensive, often guarding the sense-cups
or sensory fields on which the sense-cups are situated, or they have a
nerve extending to them and are simply tactile in function.

The surface of the epipharynx, then, appears to be highly sensitive, and
to afford the principal seat of the gustatory organs, which are
described under the head of organs of taste.

LITERATURE ON THE EPIPHARYNX

=Réaumur.= Mémoires pour servir à l’histoire des insectes, v, 1740, p.
318, Pl. 28, Figs. 4, 7, 8, 9, 10, 11 l.

=Kirby and Spence.= Intr. to entomology, iii, 1828, p. 457.

=De Geer.= ii, 1778; v, 26, Fig. 11, M.

=Kirby and Spence.= Pl. xii, Fig. 2 K.

=Latreille.= Organisation extérieure des insectes, p. 184. (Quoted
from Kirby and Spence.)

=Savigny.= Mémoires sur les animaux sans vertèbres. Partie I^{re},
1816, p. 12.

=Walter, Alfred.= Beiträge zur Morphologie der Schmetterlinge. Erster
Theil. Zur Morphologie des Schmetterlingsmundtheile. (Jena. Zeits.,
xviii, 1885, p. 752.)

=Cheshire, F. R.= Bees and bee-keeping, i, London, 1886, p. 93.

=Joseph, Gustav.= Zur Morphologie des Geschmacksorganes bei Inseckten.
(Amtlicher Bericht der 50 Versammlung deutscher Naturforscher u.
Artzte in München. 1877, pp. 227, 228.)

=Dimmock, George.= The anatomy of the mouth-parts and of the sucking
apparatus of some Diptera, 1881. (Also in Psyche, iii, pp. 231–241,
Pl. 1, 1882.)

=Packard, A. S.= On the epipharynx of the Panorpidæ. (Psyche, 1889, v,
pp. 159–164.)

—— Notes on the epipharynx and the epipharyngeal organs of taste in
mandibulate insects. (Psyche, v, pp. 193–199, 222–228, 1889.)

=Attachment of the head to the trunk.=—The head is either firmly
supported by the broad prothoracic segment in Orthoptera, many beetles,
etc., into which it is more or less retracted, or it is free and
attached by a slender neck, easily turning on the trunk, as in
dragon-flies, flies, etc. In some insects there are several chitinous
plates, situated on an island in the membrane on the under side of the
neck; these are the “cervical sclerites” of Sharp, occurring “in
Hymenoptera, in many Coleoptera, and in Blattidæ.”

=The basal or gular region of the head.=—At the hinder part of the head
is the opening (occipital foramen) into the trunk. The cheek (gena) is
the side of the head, and to its inner wall is attached the mandibular
muscle; it thus forms the region behind the eye and over the base of the
mandibles. In the Termitidæ, where the head is broad and flat, it forms
a distinct piece on the under side of the head bounding the gulo-mental
region (Fig. 28). In the Neuroptera (Corydalus, Fig. 29, and Mantispa,
Fig. 30) it is less definitely outlined.

[Illustration:

FIG. 29.—Head of _Corydalus cornutus_, ♂: _A_, from above. _B_, from
beneath. _C_, from the side. _a. cly_, clypeus anterior; _p. cly_,
clypeus posterior; _lbr_, labrum; _md_, mandible; _mx_, base of
first maxilla; _mp_, its palpus; _m_, mentum; _sm_, submentum;
_plpr_, palpifer; _lig_, fused second maxillæ; _ant_, antenna;
_occ_, occiput.
]

[Illustration:

FIG. 28.—Head of _Termopsis angusticollis_, seen from beneath, showing
the gena and gula: _m_, mentum; _sm_, submentum; _labr_, under side
of the labrum; _x_, hypopharyngeal chitinous support.
]

All the gulo-mental region of the head appears to represent the base of
the second maxillæ, and the question hence arises whether the submentum
is not the homologue of the cardines of the first maxillæ fused, and the
mentum that of the stipites of the latter also fused together. If this
should prove to be the case, the homologies between the two pairs of
maxillæ will be still closer than before supposed. Where the gula is
differentiated, this represents the basal piece of the second maxillæ.
In Figs. 28, 29, 30, and 31, these three pieces are clearly shown to
belong to the second maxillary segment. It is evident that these pieces
or sclerites belong to the second maxillary or labial segment of the
head, as does the occiput, which may represent the tergo-pleural portion
of the segment. Miall and Denny also regarded the submentum as the basal
piece of the second maxillæ.

[Illustration:

FIG. 30.—Head of _Mantispa brunnea_, under side: _e_, eye; other
lettering as in Fig. 29.
]

[Illustration:

FIG. 31.—Head of _Limnephilus pudicus_, under side: _e_, eye; _l_,
ligula; _p_, palpifer; _lp_, labial palpi.
]

=The occiput= (Fig. 29, _B_, _C_), as stated beyond, is very rarely
present as a separate piece; in the adult insect we have only observed
it in Corydalus. The occipital region may be designated as that part of
the head adjoining and containing the occipital foramen. Newport
considers the occiput as that portion of the base of the head “which is
articulated with the anterior margin of the prothorax. It is perforated
by a large foramen, through which the organs of the head are connected
with those of the trunk. It is very distinct in Hydroüs and most
Coleoptera, and in some, the Staphylinidæ, Carabidæ, and Silphidæ is
constricted and extended backwards so as to form a complete neck.” (See
also p. 51.)

[Illustration:

FIG. 32.—Interior and upper and under surface of the head of _Hydroüs
piceus_: _d_, clypeus; _e_, labrum; _g_, maxilla; _h_, its palpus;
_i_, labium; _k_, labial palpus; _p_, sutura epicranii; _q_,
cotyloid cavity; _r_, torulus; _s_, _v_, laminæ squamosa; _t_,
laminaæ posteriores; _u_, tentorium; _w_, laminæ orbitales; _x_, os
transversum; _y_, articulating cavity for the mandible; _z_, os
hypopharyngeum.—After Newport.
]

=The tentorium.=—The walls of the head are supported or braced within by
two beams or endosternites passing inwards, and forming a solid
chitinous process or loop which extends in the cockroach downwards and
forwards from the lower edge of the occipital foramen. “In front it
gives off two long crura or props, which pass to the ginglymus, and are
reflected thence upon the inner surface of the clypeus, ascending as
high as the antennary socket, round which they form a kind of rim.”
(Miall and Denny.) The œsophagus passes upwards between its anterior
crura, the long flexor of the mandible lies on each side of the central
plate; the supraœsophageal ganglion rests on the plate above, and the
subœsophageal ganglion lies below it, the nerve cords which unite the
two passing through the circular aperture. (Miall and Denny.) In
Coleoptera (Hydroüs) it protects the nervous cord which passes under it.
(Newport, Fig. 32, _u_.)

[Illustration:

FIG. 33.—Posterior view of head of Anabrus; _t_, tentorium. Joutel
_del._
]

In Anabrus the tentorium is V-shaped, the two arms originating on each
side of the base of the clypeus next to the base of each mandible the
origin being indicated by two small foramina partly concealed externally
and passing inwards and backwards and uniting just before reaching the
posterior edge of the large occipital foramen (Fig. 33).

Palmén regards the tentorium as representing a pair of tracheæ (with
the cephalic spiracles) which have become modified for supports or
for muscular attachment, since he finds that in Ephemera the
tentorium breaks across the middle during exuviation, each half
being drawn out of the head like the chitinous lining of a tracheal
tube. This view is supported by Wheeler, who has shown that the
tentorium of Doryphora originates from five pairs of invaginations
of the longitudinal commissures, and which are anterior to those of
the second maxillary segment. “These invaginations grow inwards as
slender tubes, which anastomose in some places. Their lumina are
ultimately filled with chitin.” (Jour. Morph., iii, p. 368.)

This view has also been held by Carrière and Cholodkowsky, but
Heymons concludes from his embryological studies on Forficula and
Blattidæ (1895) that it is unfounded. That this is probably the case
is proved by the fact that the apodemes of the thoracic region are
evidently not modified tracheæ, since the stigmata and tracheæ are
present.

=Number of segments in the head.=—While it is taken for granted by many
entomologists that the head of insects represents a single segment,
despite the circumstance that it bears four pairs of appendages, the
more careful, philosophical observers have recognized the fact that it
is composed of more than a single segment. Burmeister recognized only
two segments in the head; Carus and Audouin recognized three; Macleay
and Newman four; Straus-Durckheim even so many as seven. Huxley supposed
that there are five segments bearing appendages, remarking, “if the eyes
be taken to represent the appendages of another somite, the insect head
will contain six somites.” (Manual of Anat. Invert. Animals, p. 398.)

These discordant views were based on the examination of the head in
adult insects; but if we confine ourselves to the imago alone, it is
impossible to arrive at a solution of the problem.

Newport took a step in the right direction when he wrote: “It is only by
comparing the distinctly indicated parts of the head in the perfect
insect with similar ones in the larva that we can hope to ascertain the
exact number of segments of which it is composed.” He then states that
in the head of _Hydroüs piceus_ are the remains of four segments, though
still in the next paragraph, when speaking of the head as a whole, he
considers it as the first segment, “while,” he adds, “the aggregation of
segments of which it is composed we shall designate individually
_subsegments_.”

That the head of insects is composed of four segments was shown on
embryological grounds by the writer (1871) and afterwards by Graber
(1879). The antennæ and mouth-parts are outgrowths budding out from the
four primitive segments of the head; the antennæ grow out from the under
side of the procephalic lobes, and these should therefore receive the
name of antennal lobes. In like manner the mandibles and first and
second maxillæ arise respectively from the three succeeding segments.

[Illustration:

FIG. 34.—Embryo of _Anurida maritima_: _tc. ap_, minute temporary
appendage of the tritocerebral segment, the premandibular appendage;
_at_, antenna; _md_, mandible; _mx_^1, first maxilla; _mx_^2, second
maxilla; _p_^1–_p_^3, thoracic; _ap_^1, _ap_^2, abdominal
appendages; _an_, anus—After Wheeler.
]

While the postoral segments and their appendages are readily seen to be
four in number, the question arises as to whether the eyes represent the
appendages of one or more preoral segments. In this case embryology thus
far has not afforded clear, indubitable evidence. We are therefore
obliged to rely on the number of neuromeres, or primitive ganglia. In
the postoral region of the head, as also in the trunk, a pair of
neuromeres correspond to each segment. (See also under Nervous System,
and under Embryology.) We therefore turn to the primitive number of
neuromeres constituting the procephalic lobes or brain.

From the researches of Patten, Viallanes, and of Wheeler, especially of
Viallanes, it appears that the brain or supraœsophageal ganglion is
divided into three primitive segments. (See Nervous System, Brain.) The
antennæ are innervated from the middle division or deutocerebrum. Hence
the ocular segment, _i.e._ that bearing the compound and simple eyes, is
supposed to represent the first segment of the head. This, however, does
not involve the conclusion that the eyes are the homologues of the
limbs, however it may be in the Crustacea.

The second head-segment is the antennal, the antennæ being the first
pair of true jointed appendages.

The third segment of the head is very obscurely indicated, and the facts
in proof of its existence are scanty and need farther elucidation.

Viallanes’ tritocerebral lobes or division of the brain is situated in a
segment found by Wheeler to be intercalated between the antennal and
mandibular segments. He also detected in _Anurida maritima_, the
rudiments of a pair of appendages, smaller than those next to it, and
which soon disappear (Fig. 34, _tc. ap_). He calls this segment the
intercalary.[12] Heymons (1895) designates it as the “Vorkiefersegment,”
and it may thus be termed the premandibular segment.

[Illustration:

FIG. 35.—Head of embryo of honey bee: _B_, a little later stage than
_A_. _pr.m_, premandibular segment; _cl_, clypeus; _ant_, antenna;
_md_, mandible; _mx_, first maxilla; _mx′_, second maxilla; _sp_,
spiracle.—After Bütschli.
]

As early as 1870 Bütschli observed in the embryo of the honey bee the
rudiments of what appeared to be a pair of appendages between the
antennæ and mandibles, but, judging by his figures, nearer to and more
like the mandibles than the rudimentary antennæ (Fig. 35); they seemed
to him “almost like a pair of inner antennæ.”

“I find,” he says, “in no other insects any indication of this peculiar
appendage, which at the time of its greatest development attains a
larger size than the antennæ, and which, afterwards becoming less
distinct, forms by fusion with that on the other side a sort of larval
lower lip. That this appendage does not belong to the category of
segmental appendages is indicated by the site of its origin on the upper
side of the primitive band.” (Zeitschr. wissen. Zool., xx, p. 538.)

Grassi has also observed it in Apis, and regards it as the germ of a
first, but deciduous, pair of jaws. In the embryo of Hylotoma Graber
(Figs. 134, 135) found what he calls three pairs of “preantennal
projections,” one of which he thinks corresponds to the “inner antennæ”
of Bütschli. This subject needs further investigation.

It thus appears that the procephalic lobes of the embryo of insects,
with the rudiments of the antennæ, constitute the primitive head, and
perhaps correspond to the annelidan head, while gradually the antennal
appendages were in the phylogenetic development of the class fused with
the two segments of the primary head. That the second maxillary segment,
the occiput, was the last to be added, and at first somewhat
corresponded in position to the poison-fangs of centipedes (Chilopods),
is shown by our observations on the embryology of Æschna (Fig. 36).

[Illustration:

FIG. 36.—Æschna nearly ready to hatch: 4, labium, between _T_ and _e_
the occipital tergite; 5–7, legs.
]

[Illustration:

FIG. 37.—Head of embryo Nematus, showing the labial segment: _occ_,
forming the occiput; _cl_, clypeus; _lb_, labrum; _md_, mandible;
_mdm_, muscle of same; _mx_, maxilla; _mx′_, second maxilla
(labium); _oe_, œsophagus.
]

The mandibular segment appears to form a large part of the post-antennal
region of the epicranium on account of the great mandibular muscle which
arises from so large an area of the anterior region of the head (Fig.
37).

Judging from the embryo of Nematus (Fig. 37), the first maxillary
segment is tergally aborted, there being no tergo-pleural portion
left.[13]

The second maxillary segment tergally appears to be represented by the
occipital region of the head.

All the gular region, including the submentum and mentum, probably
represents the base of the labium or second maxillæ.[14] The so-called
“occiput” forms the base of the head of Corydalus, a neuropterous
insect, which, however, is more distinct in the larva. In most other
adult insects the occiput is either obsolete or fused with the hinder
part of the epicranium. We have traced the history of this piece
(sclerite) in the embryo of Æschna, a dragon-fly, and have found that it
represents the tergal portion of the sixth or labial segment. In our
memoir on the development of this dragon-fly, Pl. 2, Fig. 9, the head of
the embryo is seen to be divided into two regions, the anterior, formed
of the antennal, mandibular, and first maxillary segments, and the
posterior, formed of the sixth or labial segment. This postoral segment
at first appears to be one of the thoracic segments, but is afterwards
added to the head, though not until after birth, as it is still separate
in the freshly hatched nymph (Fig. 4; see also Kolbe, p. 132, Fig. 59,
_sq._ 5). A. Brandt’s figure of _Calopteryx virgo_ (Pl. 2, Fig. 19)
represents an embryo of a stage similar to ours, in which the postoral
or sixth (labial) segment is quite separate from the rest of the head.
The accompanying figure, copied from our memoir, also shows in a saw-fly
larva (_Nematus ventricosus_) the relations of the labial or sixth
segment to the rest of the head. The suture between the labial segment
and the preoral part of the head disappears in adult life. From this
sketch it would seem that the back part of the head, _i.e._ of the
epicranium, may be made up in part of the tergite or pleurites of the
mandibular segment, since the mandibular muscles are inserted on the
roof of the head behind the eyes. It is this labial segment which in
Corydalus evidently forms the occiput, and of which in most other
insects there is no trace in larval or adult life, unless we except
certain Orthoptera (Locusta), and the larva of the Dyticidæ.

The following table is designed to show the number and succession of the
segments of the head, with their respective segments.

TABULAR VIEW OF THE SEGMENTS, PIECES (SCLERITES), AND APPENDAGES OF THE
HEAD

═══════════════════════════════╤═══════════════════╤═══════════════════
NAME OF SEGMENT │ PIECES OR REGIONS │ APPENDAGES, ETC.
│OF THE HEAD-CAPSULE│
───────────┬───────────────────┼───────────────────┼───────────────────
│1. Ocellar │Epicranium, │Compound and simple
_Preoral_, │ (Protocerebral). │ anterior region │ eyes (Ocelli).
in early │ │ with the clypeus │
embryo. │ │ labrum, and │
│ │ epipharynx. │
│ │ │
_Postoral_,│2. Antennal │Epicranium, │Antennæ.
in early │ (Deutocerebral). │ including the │
embryo. │ │ antennal sockets.│
│ │ │
│3. Premandibular, │Wanting in │Premandibular
│ or intercalary │ postembryonic │ appendages (in
│ (Tritocerebral). │ life, except in │ Campodea).
│ │ Campodea. │
│ │ │
│4. Mandibular. │Epicranium behind │Mandibles.
│ │ the antennæ, │
│ │ genæ. │
│ │ │
│5. 1st Maxillary. │Epicranium, hinder │1st Maxillæ.
│ │ edge? Tentorium. │
│ │ │
│6. 2d Maxillary, or│Occiput. │2d Maxillæ or
│ labial. │ │ Labium.
│ │ │ Post-gula, gula,
│ │ │ submentum,
│ │ │ mentum,
│ │ │ hypopharynx
│ │ │ (lingua, ligula),
│ │ │ paraglossæ,
│ │ │ spinneret.
───────────┴───────────────────┴───────────────────┴───────────────────

[Illustration:

FIG. 38.—Larva (_a_) of a chalcid, about to pupate, with the head,
including the eyes and three ocelli, in the prothoracic segment:
_b_, _c_, pupa.
]

=The composition of the head in the Hymenoptera.=—Ratzeburg stated in
1832 that the head in the adult Hymenoptera (Cynips, Hemiteles, and
Formica) does not correspond to that of the larva, but is derived from
the head and the first thoracic segment of the larva. Westwood and also
Goureau made less complete but similar observations, though Westwood
afterwards changed his opinion, and the same view was maintained by
Reinhard. Our own observations (as seen in Fig. 38) led us to suppose
that this was a mistaken view; that the larval head, being too small to
contain that of the semipupa, was simply pushed forward, as in
caterpillars. Bugnion, however, reaffirms it in such a detailed way that
we reproduce his account. He maintains that the views of Ratzeburg are
exact and easy to verify in the chalcid genus Encyrtus, except, however,
that which concerns the ventral part and the posterior border of the
prothoracic segment.

As the time of transformation approaches, the head of the larva, he
says, is depressed and soon concealed under the edge of the prothoracic
segment; the latter elongates, becomes thicker and more convex, and
within can be seen the two oculo-cephalic imaginal buds. The head of the
perfect insect is derived not only from the head of the larva, but also
from the portion of the prothoracic segment which is occupied by the
buds, _i.e._ almost its entire dorsolateral face. But the hinder and
ventral part of this segment (which contains the imaginal buds of the
first pair of legs) takes no part in the formation of the head; these
parts, according to Bugnion, towards the end of the larval period
detaching themselves so as to become fused with the thorax and
constitute the pronotum and the prosternum.

[Illustration:

FIG. 39.—Anterior half of larva of Encyrtus, ventral face, showing the
upper (wing) and lower (leg) thoracic imaginal buds: _b_, mouth;
_ch_, chitinous arch; _gl_, silk gland; _g_, brain; _n_, nervous
cord; _a_^1, bud of fore, _a_^2, bud of hind, wing; _p_^1–_p_^3,
buds of legs; _st_^1–_st_^3, stigmata.
]

[Illustration:

FIG. 40.—Anterior part of Encyrtus larva, 1.2 mm. in length; dorsal
face; the cellular masses beginning to form the buds of the wings,
eyes, and antennæ: _o_, eye bud; _e_, stomach.
]

[Illustration:

FIG. 41.—Older Encyrtus larva, lateral view, showing the buds of the
antennæ (_f_), legs, and wings; _oe_, œsophagus; _q_^1, _q_^2,
_q_^3, buds of the genital armature; _x_, rudiment of the sexual
gland (ovary or testis); _u_, urinary tube; _i_, intestine (rectum);
_a_, anus.
]

[Illustration:

FIG. 42.—A still older larva, ready to transform. The imaginal buds of
the antennæ, eyes, wings, and legs have become elongated; lettering
as in Fig. 41.—This and Figs. 39–41 after Bugnion.
]

This mode of formation of the head may be observed still more easily
in Rhodites, Hemiteles, and Microgaster, from the fact that their
oculo-cephalic buds are much more precocious, and that the eyes are
charged with pigment at a period when the insect still preserves its
larval form.

“... I believe that this mode of formation of the head occurs in all
Hymenoptera with apodous larvæ, in this sense; that a more or less
considerable part of the first thoracic segment is always soldered
to the head of the larva to constitute the head of the perfect
insect. The arrangement of the nervous system is naturally in accord
with this peculiarity of development, and the cephalic ganglia of
the larva to which the ocular blastems later adapt themselves, are
found not in the head, but in the succeeding segment (Figs. 39, 40,
41).

“Relying on these facts, I maintain that the encroachment of the
head on the prothorax is a consequence of the preponderance in size
of the brain, and indicates the superiority of the Hymenoptera over
other insects....”

That the pronotum is derived from the larval prothoracic segment is
proved by the fact that the first pair of stigmata becomes what
authors call the “prothoracic” stigmata of the perfect insect. But
Bugnion thinks that the projection which carries it, and which he
calls the shoulder (Figs. 41 and 42), belongs to the mesonotum.

_b._ Appendages of the head

=The antennæ.=—These are organs of tactile sense, but also bear
olfactory, and in some cases auditory organs; they are usually inserted
between or in front of the eyes, and moved by two small muscles at the
base, within the head. In the more generalized insects the antennæ are
simple, many-jointed appendages, the joints being equal in size and
shape. The antennæ articulate with the head by a ball and socket joint,
the part on which it moves being called the _torulus_ (Fig. 32, _r_). In
the more specialized forms it is divided into the _scape_, the
_pedicel_, and a _flagellum_ (or _clavola_); but usually, as in ants,
wasps, and bees, there are two parts, the basal three-jointed one being
the scape, and the distal one, the usually long filiform flagellum. The
antennæ, especially the flagellum, vary greatly in form in insects of
different families and orders, this variation being the result of
adaptation to their peculiar surroundings and habits. The number of
antennal joints may be one (Articerus, a clavigerid beetle), or two in
Paussus and in _Adranes cœcus_ (Fig. 43^{12}), where they are short and
club-shaped; in flies (Muscidæ, etc.), they are very short and with few
joints, and when at rest lying in a cavity adapted for their reception.
In the lamellicorn beetles the flagellum is divided into several leaves,
and this condition may be approached in the serrate or flabellicorn
antennæ of other beetles. In Lepidoptera, and in certain saw-flies and
beetles, they are either pectinate or bipectinate, being in one case at
least, that of the Australian Hepialid (_Abantiades argenteus_),
tripectinate (Fig. 44), and in the dipterous (Tachinid) genus Talarocera
the third joint is bipectinate (Fig. 45). In Xenos and in Parnus they
may be deeply forked, while in Otiocerus, two long processes arise from
the base, giving it a trifid shape. In dragon-flies and cicadæ, they are
minute and hair-like, though jointed, while in the larvæ of many
metabolous insects they are reduced to minute three-jointed tubercles.
In aquatic beetles, bugs, etc., the antennæ are short, and often, when
at rest, bent close to the body, as long antennæ would impede their
progress.

[Illustration:

FIG. 43.—Different forms of antennæ of beetles: 1, serrate; 2,
pectinate; 3, capitate (and also geniculate); 4–7, clavate; 8, 9,
lamellate; 10, serrate (Dorcatoma); 11, irregular (Gyrinus); 12,
two-jointed antenna of _Adranes cæcus_.—After LeConte. _a_, first
joint of flagellum of antenna of _Troctes silvarum_; _b_, of _T.
divinatorius_.—After Kolbe.
]

[Illustration:

FIG. 44.—Tripectinate antenna of an Australian moth.
]

While usually more or less sensorial in function, Graber states that the
longicorn beetles in walking along a slender twig use their antennæ as a
rope-dancer does his balancing pole.

[Illustration:

FIG. 45.—Antenna of _Talarocera nigripennis_, ♂.—After Williston.
]

Recent examination of the sense-organs in the antennæ of an ant, wasp,
or bee enables us, he says, to realize what wonderful organs the antennæ
are. In such insects we have a rod-like tube which can be folded up or
extended out into space, containing the antennal nerve, which arises
directly from the brain and sends a branch to each of the thousands of
olfactory pits or pegs which stud its surface. The antenna is thus a
wonderfully complex organ, and the insect must be far more sensitive to
movements of the air, to odors, wave-sounds, and light-waves, than any
of the vertebrate animals.

That ants appear to communicate with each other, apparently talking with
their antennæ, shows the highly sensitive nature of these appendages.
“The honey-bee when constructing its cells ascertains their proper
direction and size by means of the extremities of these organs.”
(Newport.)

How dependent insects are upon their antennæ is seen when we cut them
off. The insect is at once seriously affected, its central nervous
system receiving a great shock, while it gives no such sign of distress
and loss of mental power when we remove the palpi or legs. On depriving
a bee of its antennæ, it falls helpless and partially paralyzed to the
earth, is unable at first to walk, but on partly recovering the use of
its limbs, it still has lost the power of coördinating its movements,
nor can it sting; in a few minutes, however, it becomes able to feebly
walk a few steps, but it remains over an hour nearly motionless. Other
insects after similar treatment are not so deeply affected, though bees,
wasps, ants, moths, certain beetles, and dragon-flies are at first more
or less stunned and confused.

The antennæ afford salient secondary sexual differences, as seen in the
broadly pectinated antennæ of male bombycine moths, certain saw-flies
(Lophyrus), and many other insects.

The mouth-parts, buccal appendages, or trophi, comprise, besides the
labrum, the mandibles and maxillæ.

=The mandibles.=—These are true jaws, adapted for cutting, tearing, or
crushing the food, or for defence, while in the bees they are used as
tools for modelling in wax, and in Cetonia, etc., as a brush for
collecting pollen. They are usually opposed to each other at the tips,
but in many carnivorous forms their tips cross each other like shears.
They are situated below the clypeus on each side, and are hinged to the
head by a true ginglymus articulation, consisting of two condyles or
tubercles to which muscles are attached, the principal ones being the
flexor and great extensor (Fig. 48). They are solid, chitinous, of
varied shapes, and in the form of the teeth those of the same pair
differ somewhat from each other (Fig. 46 _A_). In the pollen-eating
beetles (Cetoniæ) and in the dung-beetles (Aphodius, etc.) the edge is
soft and flexible. In the males of Lucanus, etc. (Fig. 47), and of
Corydalus (Fig. 29), they are of colossal size, and are large and
sabre-shaped in the larvæ of water-beetles, ant-lions, Chrysopa, etc.
where they are perforated at the tips, through which the blood of their
prey is sucked.

While the mandibles are generally regarded as composed of a single
piece, in Campodea and Machilis there appears to be an additional basal
piece apparently corresponding to the stipes of the first maxilla, and
separated by a faint suture from the molar or distal joint. In Campodea
there is a minute movable appendage figured both by Meinert and by
Nassonow, which appears to represent the lacinia of the maxilla (Fig.
48). Wood-Mason has observed in the mandibles of the embryo of a
Javanese cockroach, _Blatta (Panesthia) javanica_, indications of “the
same number of joints as in that of chilognathous myriopods, or one less
than in that of Machilis.” Also he adds: “In both ‘larvæ’ and adults of
_Panesthia javanica_ a faint groove crosses the ‘back’ of the mandible
at the base. This groove appears to be the remains of the joint between
the third and apical segments of the formerly 4–segmented mandibles.”

[Illustration:

FIG. 46.—Various forms of mandibles. _A_, right and left of Termopsis.
_A′_, showing at the shaded portion the “molar” of Smith. _B_,
_Termes flavipes_, soldier; _md_, its mandible. _C_, Panorpa.
]

[Illustration:

FIG. 47.—_Chiasognathus grantii_, reduced. Male.—After Darwin.
]

[Illustration:

FIG. 48.—Mandible of Campodea: _l_, prostheca or lacinia; _g_, galea;
_f_, _f_, flexor muscles; _e_, extensor; _r_, _r_, retractor; _rt_,
muscle retaining the mandible in its place.—After Meinert. _A_,
extremity of the same.—After Nassonow.
]

[Illustration:

FIG. 49.—Mandible of _Passalus cornutus_ with the prostheca (_l_):
_A_, that of a Nicaraguan species; _a_, inside, _b_, outside view,
with the muscle.
]

He also refers to the _prostheca_ of Kirby and Spence (Fig. 49), which
he thinks appears to be a mandibular lacinia homologous with it in
Staphylinidæ and other beetles (J. B. Smith also considers it as
“homologous to the lacinia of the maxilla”), and on examining it in _P.
cornutus_ and a Nicaragua species (Fig. 49), we adopt his view, since we
have found that it is freely movable and attached by a tendon and muscle
to the galea. In the rove beetles (Goërius, Staphylinus, etc.) and in
the subaquatic Heteroceridæ, instead of a molar process, is a membranous
setose appendage not unlike the coxal appendages of Scolopendrella,
movably articulated to the jaw, which he thinks answers to the molar
branch of the jaws in Blatta and Machilis. “It has its homologue in the
diminutive Trichopterygidæ in the firmly chitinized quadrant-shaped
second mandibular joint, which is used in a peculiar manner in crushing
the food”; also in the movable tooth of the Passalidæ, and in the
membranous inner lobe of the mandibles of the goliath-beetles, etc.

J. B. Smith has clearly shown that the mandibles are compound in certain
of the lamellicorns. In _Copris carolina_ (Fig. 50), he says, the small
membranous mandibles are divided into a basal piece (basalis), the
homologue of the stipes in the maxilla; another of the basal pieces he
calls the molar, and this is the equivalent of the subgalea, while a
third sclerite, only observed in Copris, is the _conjunctivus_, the
lacinia (prostheca) being well developed. Smith therefore concludes
“that the structure of the mandible is fundamentally the same as that of
the labium and maxilla, and that we have an equally complex organ in
point of origin. Its usual function, however, demands a powerful and
solid structure, and the sclerites are in most instances as thoroughly
chitinized and so closely united to the others that practically there is
only a single piece, in which the homology is obscured.” (Trans. Amer.
Ent. Soc., xix, pp. 84, 85. 1892.) From the studies of Smith and our
observations on Staphylinus, Passalus, Phanæus, etc. (Fig. 50, _A_, _B_)
we fully agree with the view that the mandibles are primarily 3–lobed
appendages like the maxillæ. Nymphal Ephemerids have a lacinia-like
process. (Heymons.)

[Illustration:

FIG. 50.—Mandible of _Copris carolina_.—After Smith. _A_′_C.
anaglypticus_. _A_ (figure to right), do. of _Leistotrophus
cingulatus_; _B_, of _Phanæus carnifex_; _g′_, end of galea,—_g_,
enlarged; _c_, conjunctivus. _C_, of _Meloë angusticollis_: _l_,
lacinia; _a_, lacinia enlarged.
]

Mandibles are wanting in the adults of the more specialized Lepidoptera,
being vestigial in the most generalized forms (certain Tineina and
Crambus), but well developed in that very primitive moth, Eriocephala
(Fig. 51). They are also completely atrophied in the adult Trichoptera,
though very large and functional in the pupa of these insects (Fig. 52),
as also in the pupa of Micropteryx (Fig. 53). They are also wanting in
the imago of male Diptera and in the females of all flies except
Culicidæ and Tabanidæ.

They are said by Dr. Horn to be absent in the adult _Platypsyllus
castoris_, though well developed in the larva; and functional mandibles
are lacking in the Hemiptera.

=The first maxillæ.=—These highly differentiated appendages are inserted
on the sides of the head just behind the mandibles and the mouth, and
are divided into three lobes, or divisions, which are supported upon
two, and sometimes three basal pieces, _i.e._ the basal joint or
_cardo_, the second joint or _stipes_, with the _palpifer_, the latter
present in Termitidæ (Fig. 54, _plpgr_), but not always separately
developed (Fig. 55). The cardo varies in shape, but is more or less
triangular and is usually wedged in between the submentum and mandible.
It is succeeded by the stipes, which usually forms the support for the
three lobes of the maxilla, and is more or less square in shape.

[Illustration:

FIG. 51.—Mandible of _Eriocephala calthella_: _a_, _a′_, inner and
outer articulation; _s_, cavity of the joint (acetabulum); _A_, end
seen from one side of the cutting edge.—After Walter.
]

[Illustration:

FIG. 52.—_A_, Pupa of _Phryganea pilosa_.—After Pictet. _B_, mandibles
of pupa of _Molanna angustata_.—After Sharp.
]

[Illustration:

FIG. 53.—Pupa of _Micropteryx purpuriella_, front view: _md_,
mandibles; _mx.p_, maxillary palpus, end drawn separately; _mx.’p_,
labial palpi; _lb_, labrum; _A_, another view from a cast skin.
]

The three distal divisions of the maxilla are called, respectively,
beginning with the innermost, the _lacinia_, _galea_, and _palpifer_,
the latter being a lobe or segment bearing the _palpus_. The lacinia is
more or less jaw-like and armed on the inner edge with either flexible
or stiff bristles, spines, or teeth, which are very variable in shape
and are of use as stiff brushes in pollen-eating beetles, etc. The galea
is either single-jointed and helmet-shaped or subspatulate, as in most
Orthoptera, or 2–jointed in Gryllotalpa, or lacinia-like in Myrmeleon
(Fig. 55, _C_); or, in the Carabidæ (Fig. 56) and Cicindelidæ, it is
2–jointed and in form and function like a palpus.

[Illustration:

FIG. 54.—_A_, maxilla of _Termopsis angusticollis_. _B_, _Termes
flavipes_: _c_, cardo; _sti_, stipes; _plpgr_, palpiger; _palp_,
palpus; _lac_, lacinia; _g_, _gal_, galea.
]

[Illustration:

FIG. 55.—_A_, maxilla of _Mantispa brunnea_. _B_, _Ascalaphus
longicornis_. _C_, _Myrmeleon diversum_. Lettering as in Fig. 54.
]

[Illustration:

FIG. 56.—Maxilla of a carabid, _Anophthalmus tellkampfii_: _l_,
lacinia; _g_, 2–jointed galea; _p_, palpus; _st_, stipes; _c_,
cardo.
]

[Illustration:

FIG. 57.—Maxilla of _Nemognatha_, ♀, from Montana. _A_, base of
maxilla enlarged to show the taste-papillæ (_tp_) and cups (_tc_),
on the galea (_ga_). _B_, part of end of galea to show the imperfect
segments and taste-organs: _n_, nerve; a ganglionated nerve supplies
each taste-papilla or cup; _l_, lacinia; _p_, palpifer; _s_,
subgalea.
]

[Illustration:

FIG. 58.—Maxilla of Panorpa.
]

[Illustration:

FIG. 59.—Maxilla of _Limnephilus pudicus_: _mx_, stipes; _lac_, galea.
]

The palpus is in general antenniform and is composed of from 1 to 6
joints, being usually 4– or 5–jointed, and is much longer than the
galea. In the maxilla of the beetle Nemognatha (Fig. 57), the galea is
greatly elongated, the two together forming an imperfect tube or
proboscis and reminding one of the tongue of a moth, while the lacinia
is reduced. In the Mecoptera the lacinia and galea are closely similar
(Fig. 58); in the Trichoptera only one of the lobes is present (Fig.
59), while in the Lepidoptera the galea unites with its mate to form the
so-called tongue (Fig. 60). The maxilla of the male of _Tegeticula
yuccasella_ is normal, though the galeæ are separate; but in the female,
what Smith regards as the palpifer (the “tentacle” of Riley) is
remarkably developed, being nearly as long as the galea (Fig. 61) and
armed with stout setæ, the pair of processes being adapted for holding a
large mass of pollen under the head.

[Illustration:

FIG. 60.—Tongue of _Aletia xylina_, with the end magnified.—Pergande
_del._, from Riley. _A_, much reduced maxilla (_mx_) of _Paleacrita
vernata_; _mx.p_, palpus.
]

[Illustration:

FIG. 61.—_A_, maxilla of _Tegeticula yuccasella_, ♂: _g_, galea. _B_,
♀: _pl_, enormously developed palpifer; _mx.p_, palpus; _c_, cardo;
_st_, stipes; _sty_, stylus.
]

In coleopterous larvæ the maxillæ are 2–lobed (Fig. 62), the galea being
undifferentiated, but in those of saw-flies the galea is present (Fig.
63, _gal_).

[Illustration:

FIG. 62.—Larva of _Rhagium lineatum_: _lat_, lateral view of head and
thoracic segments; _mx_, first maxilla; _ml_, undifferentiated
lacinia and galea; _v_, under side of head and pro- and meso-
thoracic segments; _v.m.s._, one of the middle ventral segments,
magnified six times; _mx′_, 2d maxilla.
]

It now seems most probable that in the first maxillæ we have the primary
form of buccal appendage of insects, the appendage being composed of
three basal pieces with three variously modified distal lobes or
divisions; and that the mandibles and second maxillæ are modifications
of this type.

How wonderfully the maxillæ of the Lepidoptera are modified, and the
peculiar shapes assumed in the Diptera, Hymenoptera, and other groups,
will be stated in the accounts of those orders, but it is well to recall
the fact that in the most primitive and generalized moth, Eriocephala,
the lacinia is well developed (Fig. 64).

As Newport remarks, the office of the maxillæ in the mandibulate insects
is of a twofold kind; since they are adapted not only for seizing and
retaining the food in the mouth, but also as accessory jaws, since they
aid the mandibles in comminuting it before it is passed on to the
pharynx and swallowed. Hence, as the food varies so much in nature and
situation, it will be readily seen that the maxillæ, especially their
distal parts, vary correspondingly. Thus far no close observations on
the exact use of the first and second maxillæ have been published.

The palpi also are not only organs of touch, but in some cases act as
hands and also bear minute sense-organs, the function of which is
unknown, but would appear to be usually that of smell.

[Illustration:

FIG. 63.—_Selandria_ larva, common on _Carya porcina_, with details of
mouth-parts: _leg_, leg; _mx_, maxilla; _gal_, galea; _lac_,
lacinia.
]

=The second maxillæ.=—The “under-lip” or labium of insects is formed by
the fusion at the basal portion of what in the embryo are separate
appendages, and which arise in the same manner as the first maxillæ.
They are invariably solidly united, no cases of partial or incomplete
fusion being known. The so-called labium is situated in front of the
gula or gular region, and is bounded on each side by the gena, or cheek.
As already observed, the second maxillæ appear to be the appendages of
the last or occipital segment of the head.

[Illustration:

FIG. 64.—Maxilla of _Eriocephala calthella_: _l_, lacinia; _g_, galea;
_mx.p_, maxillary palpus; _st_, stipes; _c_, cardo.—After Walter.
]

The second maxillæ are very much differentiated and vary greatly in the
different orders, being especially modified in the haustellate or
suctorial orders, notably the Hymenoptera and Diptera. In the
mandibulate orders, particularly the Orthoptera, where they are most
generalized and primitive in shape and structure, they consist of the
following parts: the _gula_ (a _postgula_ is present in Dermaptera),
_submentum_ (lora of Cheshire, i, p. 91), _mentum_, _palpifer_, the
latter bearing the _palpi_; the _lingua_ (_ligula_) and _paraglossæ_,
while the hypopharynx or lingua is situated on the upper side. The
labial palpi are of the same general shape as those of the first
maxillæ, but shorter, with very rarely more than three joints, though in
Pteronarcys there are four. Leon has detected vestigial labial palpi in
several Hemiptera (Fig. 73). As to the exact nature and limits of the
gula, we are not certain; it is not always present, and may be only a
differentiation of the submentum, or the latter piece may be regarded as
a part of the gula.

We are disposed to consider the second maxillæ as morphologically nearly
the exact equivalents of the first pair of maxillæ, and if we adopt this
view it will greatly simplify our conception of the real nature of this
complicated organ. The object of the fusion of the basal portion appears
to be to form an under-lip, in order both to prevent the food from
falling backwards out of the mouth, and, with the aid of the first pair
of maxillæ, to pass it forward to be crushed between the mandibles, the
two sets of appendages acting somewhat as the tongue of vertebrates to
carry and arrange or press the morsels of food between the teeth or
cutting edges of the mandibles.

The spines often present on the free inner edges of the first and second
maxillæ (Figs. 54, 62) form rude combs which seem to clean the antennæ,
etc., often aiding the tibial combs in this operation.

The submentum and mentum, or the mentum when no submentum is
differentiated (with the gula, when present), appear to be collectively
homologous with the cardines of the first pair of maxillæ, together with
the palpifers and the stipites.[15] These pieces are more or less
square, and have a slightly marked median suture in Termitidæ, the sign
of primitive fusion or coalescence.

The most primitive form of the second maxillæ occurs in the Orthoptera
and in the Termitidæ. The palpifer is either single (Periplaneta,
Diapheromera, Gryllidæ) or double (_Blatta orientalis_, Locustidæ). In
Prisopus the single piece in front of the palpifer is in other forms
divided, each half (Blatta, Locustidæ, Acrydidæ) bearing the two
“paraglossæ,” which appendages in reality are the homologues of the
lacinia and galea of the first maxillæ.[16] In the Termitidæ (Fig. 65)
the lingua is not differentiated from the palpifer, and the two
paraglossæ (or the lamina externa and interna of some authors) with the
palpus are easily seen to be the homologues of the three lobes of the
first maxillæ. In the Perlidæ (Pteronarcys, Fig. 66) the palpifer is
divided, while the four paraglossæ arise, as in Prisopus and
Anisomorpha, from an undivided piece, the lingua not being visible from
without. In the Neuroptera the lingua or ligula is a large, broad,
single lobe, without “paraglossæ,” and the palpifer is either single
(Myrmeleon, Fig. 67), or divided (Mantispa, Fig. 68). In Corydalus (Fig.
29) the palpifer forms a single piece, and the lingua is undivided,
though lobed on the free edge.

[Illustration:

FIG. 65.—Second maxillæ of _Termopsis angusticollis_: _li_, the
homologue of the lacinia; _le_, galea.
]

In the metabolic orders above the Neuroptera the lingua is variously
modified, or specialized, with no vestiges of the lacinia or galea,
except in that very primitive moth, Eriocephala, in which Walter found a
minute free galea, _me_, and an inner lobe (Figs. 76, 77), the lacinia.

[Illustration:

FIG. 66.—Second maxillæ of _Pteronarcys californica_.
]

[Illustration:

FIG. 67.—Second maxillæ of _Myrmeleon diversum_.
]

[Illustration:

FIG. 68.—Second maxillæ of _Mantispa brunnea_.
]

=The hypopharynx.=—While in its most generalized condition, as in
Synaptera, Dermaptera, Orthoptera, and Neuroptera, this anterior median
fold or outgrowth of the labium forming the floor of the mouth may
retain the designation of “tongue,” lingua, or ligula; in its more
specialized form, particularly when used as a piercing or lapping organ,
the use of the name _hypopharynx_ seems most desirable. And this is
especially the case since, like the epipharynx, it is morphologically a
median structure, and while the epipharynx forms the soft, sensitive
roof of the mouth, or pharynx; its opposite, the hypopharynx, rises as a
fold from the floor of the mouth, forming in its most generalized
condition a specialized fold of the buccal integument. In certain cases,
as in the honey-bee, the very long slender “tongue” or hypopharynx is
evidently, as in the case of the epipharynx, a highly sensitive armature
of the mouth.

In all insects this organ—whether forming a soft, tongue-like, anterior
portion or fold of the labium, and “continuous with the lower wall of
the pharynx,” or a hard, piercing, awl-like appendage (fleas and flies),
or a long, slender, hairy or setose, trough-like structure like the
“tongue” of the honey-bee—has a definite location at the end and on the
upper side of the labium, and serves to receive at its base the external
opening of the salivary duct.

The hypopharynx, as well shown in its lingua condition in Orthoptera, is
continuous with and forms the anterior part or fold of the base of the
coalesced second maxillæ. It does not seem to be paired, or to represent
a pair of appendages.

Opinion regarding the homology of this unpaired piercing organ is by no
means settled, and while there is a general agreement as to the nature
of the paired mouth-parts, recent observers differ very much as to the
morphology of the organ in question.

It is the langue or _lingua_ of Savigny (1816), the _ligula_ of Kirby
and Spence (1828), the _langue ou languette_ (_lancette médiane du
suçoir_) of Dugès (1832), the _lingua_ of Westwood (Class, ins., ii, p.
489, 1840), “the unpaired median piercing organ” (“the analogon of the
epipharynx of Diptera”) of Karsten (1864), the “tongue” of Taschenberg
(1880).

The name _hypopharynx_ was first proposed by Savigny in 1816, who, after
naming the membranous plate which has for its base the upper side of the
pharynx, the _epipharynx_, remarks: “Dans quelques genres, notamment
dans les Eucères, le bord inférieur de ce même pharynx donne naissance à
un autre appendice plus solide que le précédent, et qui s’emboîte avec
lui. Je donnerai à ce dernier le nom de _langue_ ou d’_hypopharynx_.
Voilà donc la bouche des Hyménoptères composée de quatre organes
impaires, sans y comprendre la ganache ou le menton; savoir, la lèvre
supérieure, l’épipharynx, l’hypopharynx, et la lèvre inférieure, et de
deux organes paires, les mandibules et les mâchoires.”

As stated by Dimmock: “The hypopharynx is usually present in Diptera
(according to Menzbier absent in Sargus), and contains a tube, opening
by a channel on its upper surface; this channel extends back, more or
less, from the tip, and is the outlet for the salivary secretion. The
tip of the hypopharynx may be naked and used as a lance (Hæmatopota,
according to Menzbier), or may be hairy (Musca). The upper side of the
base of the hypopharynx is continuous with the lower wall of the
pharynx; its under surface may entirely coalesce with the labium (Culex,
male), may join the labium more or less, anterior to the month (Musca),
or, if either mandibles or maxillæ are present, its base may join them
(Culex, female).” (p. 43.)

[Illustration:

FIG. 69.—Section of head of _Machilis maritima_: _hyp_, hypopharynx;
_lbr_, labrum; _t_, tentorium; _ph_, room in which the mandibles
move on each other; _p_, paraglossa; _mx_, labium; _sd_, salivary
duct; _s.gl_, salivary gland. _oe_, œsophagus.—After Oudemans.
]

We will now briefly describe the lingua, first of the mandibulate or
biting insects, and then its specialized form, the hypopharynx of the
haustellate and lapping insects.

The lingua (hypopharynx) exists in perhaps its most generalized
condition in the Thysanura (Fig. 69), where it forms a soft projection,
having the same relations as in Anabrus and other Orthoptera.[17]

In the cockroach (Fig. 70), as stated by Miall and Denny, the lingua is
a chitinous fold of the oral integument situated in front of the labium,
and lying in the cavity of the mouth. The common duct of the salivary
glands enters the lingua, and opens on its hinder surface. The lingua is
supported by a chitinous skeleton (Figs. 70, _B_; 82, _shp_). “The thin
chitinous surface of the lingua is hairy, like other parts of the mouth,
and stiffened by special chitinous rods or bands.” (Miall and Denny.)

[Illustration:

FIG. 70.—Hypopharynx of _Periplaneta orientalis_; the arrow points out
of the opening of the salivary duct: _A_, origin of salivary duct.
_B_, side view. _C_, front view.—After Miall and Denny.
]

In the Acrydiidæ (_Melanoplus femur-rubrum_) the tongue is a large,
membranous, partly hollow expansion of the base of the labium. It may be
exposed by depressing the end of the labium, when the opening of the
salivary duct may be seen at the bottom or end of the space or gap
between the hinder base of the tongue, and the inner anterior base of
the labium, as shown by the arrows in Fig. 70. It is somewhat pyriform,
slightly keeled above, and bearing fine stiff bristles, which, as they
point more or less inwards, probably aid in retaining the food within
the mouth. The base of the tongue is narrow, and extends back to near
the pharynx, there being on the floor of the mouth, behind the tongue,
two oblique, slight ridges, covered with stiff, golden-yellow hairs,
like those on the tongue. The opening of the salivary duct is situated
on the under or hinder side of the hypopharynx, between it and the base
of the labium, the base of the former being cleft; the hollow thus
formed is situated over the opening, and forms the salivary receptacle.

[Illustration:

FIG. 71.—Section through the anterior part of the head of Anabrus (the
mandibles removed), showing the relations of the hypopharynx (_hyp_)
to the opening of the salivary duct (_sd_): _g_, galea; _l_,
lacinia; _mt_, mentum; _oe_, œsophagus; _lbr_, labrum; _cl_,
clypeus.
]

In the Locustidæ (Anabrus, Fig. 71) the tongue (hypopharynx) is a broad,
somewhat flattened lobe arising from the upper part of the base of the
mentum and behind the palpifer. This lobe is cavernous underneath, the
hollow being the salivary receptacle (_sr_); the latter is situated over
the opening of the salivary duct, which is placed between the base of
both the hypopharynx and the labium. The salivary fluid apparently has
to pass up and around on each side of the hypopharynx in order to mix
with the food.

These relations in the Orthoptera are also the same in the Perlidæ,
where the hypopharynx is well developed, forming an unusually large
tongue-like mass, nearly filling the buccal cavity.

[Illustration:

FIG. 72.—Lingua of a May-fly, _Heptagenia longicauda_, ×16: _m_,
central; _l_, lateral pieces.—After Vayssière from Sharp.
]

In the Odonata the lingua is a small, rounded lobe, as also in the
Ephemeridæ; in the nymph, however, of Heptagenia (Fig. 72) it is highly
developed, according to Vayssière, who seems inclined to regard it as
representing a pair of appendages. The tongue in Hemiptera is said by
Léon to be present in _Benacus griseus_ (Say) and to correspond to the
subgalea of Brullé or hypodactyle of Audouin (Fig. 73), but this appears
to correspond to the labium proper, rather than a true lingua, the
latter not being differentiated in this order. In the Coleoptera the
lingua is rather small. In beetles, as Anopthalmus (Fig. 74), it forms a
setose lobe; and a well-developed nerve, the lingual nerve, passes to
it, dividing at the end into several branches (_n-l_). In Sialis the
lingua is short, much less developed than usual, being rounded, and
bears on the edge what appear to be numerous taste-hairs, like those on
the ends of the maxillary and labial palpi.

[Illustration:

FIG. 73.—_A_, labium of _Zaitha anura_. _B_, of _Z. margineguttata_.
_C_, of _Gerris najas_: _mt_, mentum; _lp_, labial palpi; _sg_,
subgalea; _l_, lacinia (= intermaxillare and præmaxillare of
Brullé); _g_, galea.—After Léon.
]

In the adult Panorpidæ the lingua is a minute, simple lobe.

[Illustration:

FIG. 74.—Section through head of a carabid, _Anopthalmus telkampfii_:
_br_, brain; _f. g_, frontal ganglion; _soe_, subœsophageal
ganglion; _co_, commissure; _n. l_, nerve sending branches to the
lingua (_l_); _mn_, maxillary nerve; _mx_, 1st maxilla; _mm_,
maxillary muscle; _mx′_, 2d maxilla; _mt_, muscle of mentum; _le_,
elevator muscle of the œsophagus; _l_ of the clypeus, and a third
beyond raising the labrum (_lbr_); _eph_, epipharynx; _g_, _g_,
salivary glands above; _g_^2, lingual gland below the œsophagus
(_oe_); _m_, mouth; _pv_, proventriculus; _md_, mandible. _A_,
section passing through lingual gland (_g_^2).
]

In the larval Trichoptera the spinneret is well developed, and in
structure substantially like that of caterpillars, and it is plainly the
homologue of the hypopharynx, receiving as it does the end of the
silk-duct.

In the adult Trichoptera the hypopharynx is a very large, tongue-like,
fleshy outgrowth, and is, both in situation and structure, since it
contains the opening of the silk-duct, exactly homologous with the
hypopharynx of insects of other orders, being somewhat intermediate
between the fleshy tongue or lingua of the mandibulate insects,
especially the Neuroptera, and the hypopharynx of the bees (Fig. 86).
Lucas describes and figures it under the name of “haustellum,” but does
not homologize it with the hypopharynx. The caddis-flies have been
observed to drink water and take in both fluid and fine particles of
solid food, and to use the haustellum for this purpose, the end being
provided with minute sense-organs like those on the first maxillary
lacinia, and possibly of a gustatory nature.

[Illustration:

FIG. 75.—Head of _Anabolia furcata_: _A_, front view, showing the
labrum removed. _B_, side view; _ant_, antenna; _oc_, ocellus; _ol_,
labrum; _gh_, articulatory process; _cmx__{1}, cardo; _stmx__{1},
stipes; _lemx__{1}, outer lobe (galea); _ptmx__{1}, palpus of 1st
maxilla; _pl_, palpus of 2d maxilla; _ha_, haustellum; _so_,
gustatory pits; _spr_, opening of salivary duct; _chsp_, chitinous
hook of the clasp; _spr_, furrow or gutter of the haustellum.—After
Lucas.
]

[Illustration:

FIG. 76.—Hypopharynx of _Eriocephala calthella_: _lig_, ligula, its
membranous hinder edge; _lig′_, anterior horny edge of the
ligula-tube opening outwards; _hp_, contour of the hypopharynx;
_mi_, mala interior (lacinia); _me_, mala exterior (galea), of
second maxilla; _mx′ p_, labial palpus.—After Walter.
]

The spinneret of the larvæ of Lepidoptera is evidently the homologue of
the hypopharynx of insects of other orders. It will be seen that the
homology of the different parts is identical, the common duct of the
silk-glands opening at the end of the hypopharynx, which here forms a
complete tube or proboscis extending beyond the end of the labium, in
adaptation to its use as a spinning organ.

Walter refers to Burgess’s discovery of a hypopharynx in _Danais
archippus_, remarking that this organ in the adult Eriocephalidæ (Fig.
76) exhibits a great similarity to the relations observable in the lower
insects, adding:—

[Illustration:

FIG. 77.— Labium of _Micropteryx anderschella_ seen from within (the
labial palpi (_mx.′ p_) removed to their basal joint). Lettering as
in Fig. 76.—After Walter.
]

“The furrow is here within coalesced with the inner side of the labium,
and though I see in the entire structure of the head the inner edge of
the ligula tube extended under the epipharynx as far as the mandible, I
must also accept the fact that here also the hypopharynx extends to the
mouth-opening as in all other sucking insects with a well-developed
under-lip, viz. the Diptera and Hymenoptera.”

He has also discovered in Micropteryx a paired structure which he
regards as the hypopharynx (Fig. 77). As he states:

[Illustration:

FIG. 78.—Hypopharynx (_hph_) of Danais: _cl_, clypeus; _sd_, salivary
duct; _m_, labial palp muscles; _fm_, frontal muscle; _ph_, pharynx;
_cor_, cornea.—After Burgess.
]

“A portion of the inner surface of the tube-like ligula is covered by a
furrow-like band which, close to the inner side, is coalesced with it,
and in position, shape, as well as its appendages or teeth on the edge,
may be regarded as nothing else than the hypopharynx.”

A hypopharynx is also present in the highest Lepidoptera, Burgess having
detected it in _Danais archippus_. He states that the hypopharynx forms
the floor of the pharyngeal cavity; “it is convex on each side of a
median furrow (Fig. 78, _hph_) and somewhat resembles in shape the human
breast. The convex areas are dotted over with little papillæ, which
possibly may be taste-organs.”

As a piercing organ the hypopharynx reaches its greatest development in
the Siphonaptera and Diptera, where the chitinous parts are greatly
hypertrophied, the fleshy tongue-like portion so developed in the
mandibulate orders being greatly reduced. The chitinous parts are alike
on each side of the median organ, being bilaterally symmetrical.

[Illustration:

FIG. 79.—_A_, hypopharynx of _Pulex canis_: _x_, basal portion
situated within the head; _s. d_, common duct of the four
bladder-shaped salivary glands; _s. d′_, opening of the tubular
salivary glands into the throat. _B_, end of the hypopharynx,
showing the gutter-like structure and teeth at the end.—After
Landois.
]

[Illustration:

FIG. 80.—Beak of Vermipsylla: _hyp_, hypopharynx.—After Wagner.
]

In the fleas the hypopharynx is a large, slender, unpaired, long,
chitinous trough, as long as the mandibles, and toothed at the end.
Figures 79 and 80 show its relations to the other parts of the mouth; in
Fig. 79, _x_, is seen where the salivary duct opens into the pharynx.
Although this organ is not unanimously referred to the hypopharynx, yet
from the description of Landois and others, it is evident that this
structure does not correspond to the labrum or epipharynx, but belongs
to or arises from the floor of the mouth, and, being in close relation
to the labium, and also receiving the salivary duct, must be a true
hypopharynx.

In the Diptera the hypopharynx reaches its highest development as a
large, stout, awl-like structure.

Meinert, in his detailed and elaborately illustrated work, Trophi
Dipterorum (1881), has made an advance on our knowledge of the
hypopharynx and its homologies, both by his evidently faithful
descriptions and dissections, and by his admirably clear figures.

[Illustration:

FIG. 81.—_Culex pipiens_, section of head: _oe_ œsophagus; _sm_, upper
muscle, _lm_, lower muscle of the œsophagus; _ph_, pharynx; _rm_,
retractor muscle of the receptacle (_r_) of the salivary duct
(_s.d_); _lbr_, labrum; _ep_, left style of the epipharynx; _f_,
part of front of head.—After Meinert.
]

[Illustration:

FIG. 82.—Pharynx and hypopharynx of _Simulium fuscipes_: _lph_, lower
lamina of the pharynx; _p_, the salivary duct (_s.d_) perforating
the pharynx; _o_, orifice of the duct; _shp_, styles of the
hypopharynx; _mph_, membranous edge of the hypopharynx; _m_,
protractor muscle of the pharynx; _gp_, gustatory papillæ.—After
Meinert.
]

“The hypopharynx, a continuation of the lower edge (_lamina_) of the
pharynx, most generally free, more or less produced, acute
anteriorly, forms with the labrum the tube of the pump (_antliæ_).
(The hypopharynx when obsolete, or coalesced with the canal of the
proboscis, is the _theca_; in such a case the siphon or tube is
formed by the theca and labrum.) Meanwhile the hypopharynx, the
largest of all the trophi (_omnium trophorum maximus_), constitutes
the chief piercing organ (_telum_) of Diptera. The hypopharynx is
moved by protractor, most generally quite or very powerful, and by
retractor muscles.

“The efferent duct of the thoracic salivary glands (_ductus
salivalis_) perforates the hypopharynx, more or less near the base,
that the saliva may be ejected through the canal into the wound, or
that it may be conducted along the labella. Very rarely the salivary
duct, perforating the hypopharynx, is continued in the shape of a
free, very slender tube.

“The salivary duct behind the base of the hypopharynx forms the
receptacle or _receptaculum_, provided with retractor and levator
muscles.”

[Illustration:

FIG. 83.—Labrum-epipharynx (_lbr_ and _eph_) and hypopharynx (_hyp_)
of _Tabanus brominus_: _oe_, posterior cylindrical portion of the
œsophagus; _a_, anterior swollen portion of the same; _ph_, pharynx;
_ph.m_, pharyngeal muscle; _p.ph_, protractor muscle of the pharynx;
_r.oe_, retractor muscle of the œsophagus; _r.ph_, retractor muscle
of the pharynx; _f.oe_, flexor muscle of the pharynx; _t.oe_,
twisting muscle of the œsophagus; _s.r_, receptacle of the salivary
duct; _l_, its elevator muscle; _s_, its retractor muscle; _cl_,
clypeus.—After Meinert.
]

[Illustration:

FIG. 84.—Œsophagus (_oe_), pharynx (_ph_) with epipharynx and labrum
(_lbr_) of _Asilus atricapillus_: _m_, _ph_, pharyngeal muscle;
_sr_, salivary receptacle; _t_, twisting; _r_, _l′r_, retractor
muscles; other lettering as in Fig. 83.—After Meinert.
]

It has been carefully studied by Meinert in a species of Culex (Fig.
81), Simulium (Fig. 82), Tabanus (Fig. 83), and in Asilus (Fig. 84),
where it is seen to attain enormous proportions. In the Hymenoptera,
this organ in its most specialized condition is a trough-like rod,
adapted for lapping nectar (Fig. 85, 86, _hyp_). The tongue or
hypopharynx of the honey-bee has been elaborately described by Cheshire
in his Bees and Bee Keeping.[18] He calls it the tongue or ligula. It is
situated in a tube formed by the maxillæ and labial palpi, and can be
partially retracted into the mentum. He states that it can move up and
down in the tube thus formed, and then describes it as covered by a
hairy sheath, its great elasticity being due to a rod running through
its centre enabling it to be used as a lapping tongue. The sheath

“passes round the tongue to the back, where its edges do not meet,
but are continuous with a very thin plaited membrane (_G_, _pm_)
covered with minute hairs. This membrane, after passing towards the
sides of the tongue, returns to the angle of the nucleus, or rod,
over the under surface of which it is probably continued. The rod
passes through the tongue from end to end, gradually tapering
towards its extremity, and is best studied in the queen, where I
trace many nerve threads and cells. It is undoubtedly endowed with
voluntary movement, and must be partly muscular, although I have
failed completely in getting any evidence of striation. The rod on
the underside has a gutter, or trough-like hollow (_cd_, the central
duct) which is formed into a pseudotube (false tube) by
intercrossing of black hairs. It will also be seen that, by the
posterior meeting of the sheath, the space between the folded
membrane (_G_, _sd_) becomes two pseudotubes of larger size, which I
shall call the side ducts.

[Illustration:

FIG. 85.—Head of honey bee, worker: _a_, antenna; _g_, epipharynx;
_m_, mandible; _mx_, maxilla; _mxp_, maxillary palpus; _pg_,
paraglossa; _lp_, labial palpus; _l_, hypopharynx; _b_, its
spoon.—After Cheshire; from Bull. Div. Ent. U. S. Dept. Agr.
]

“These central and side ducts run down to that part of the tongue
where the spoon, or bouton (_K_, Fig. 86) is placed. This is
provided with very delicate split hairs (_b_, Fig. 86) capable of
brushing up the most minute quantity of nectar, which by capillarity
is at once transferred by the gathering hairs (which are here
numerous, long, and thin) to two side groove-like forms at the back
of the bouton, and which are really the opened-out extremity of the
centre and side ducts, assuming, immediately above the bouton, the
form seen in _F_, Fig. 86. The central duct, which is only from
1⁄600 inch to 1⁄1000 inch in diameter, because of its smaller size,
and so greater capillary attraction, receives the nectar, if
insufficient in quantity to fill the side ducts. But good
honey-yielding plants would bring both centre and side ducts into
requisition. The nectar is sucked up until it reaches the paraglossæ
(_pa_, _B_, Fig. 86), which are plate-like in front, but membranous
extensions, like small aprons, behind; and by these the nectar
reaches the front of the tongue, to be swallowed as before
described.”

[Illustration:

FIG. 86.—Tongue or ligula of the honey bee: _A_, under side of the
tongue; _lp_, labial palpi; _r_, _r_, rod; _p_, pouch; _sh_, sheath;
_gh_, gathering hairs; _b_, bouton or spoon. _B_, under lip or
labium, with appendages, partly dissected; _l_, lora or submentum;
_a_, _a_, retractor linguæ longus; _sd_, salivary duct; _rb_ and
_b_, retractor linguæ biceps; _mx_, maxillæ; _lp_, labial palpi;
_pa_, paraglossa; _gr_, feeding groove; _sh_, sheath of ligula. _C_,
_D_, _E_, sections of ligula; _hp_, hyaline plate of maxilla; _h_,
hairs acting as stops; _mx_, maxilla; _lp_, labial palpi; _sd_, side
duct. _F_, cross-section of extremity of tongue near the “spoon”;
_th_, tactile hairs; _r_, rod; _n_, nucleus; _gh_, gathering hairs.
_G_, cross-section of tongue without gathering hairs, × 400 times;
_sh_, sheath; _b_, blood space; _t_, trachea: _ng_, gustatory nerve;
_cd_, central duct; _sd_, lateral duct; _pm_, plaited membrane. _H_,
same as _G_, but magnified two hundred times, and with _pm_, plaited
membrane, turned outwards; _h_, closing hairs; _lp_, labial palpi;
_b_, blood; _n_, nucleus; _r_, rod; _h_, closing hairs. _I_, small
portion of the sheath; lettering as before. _K_, extremity of the
tongue, with spoon; _b_, branching hairs for gathering.—After
Cheshire.
]

Cheshire then settles the question which has been in dispute since the
time of Swammerdam, whether the bee’s tongue is solid or tubular. He
agrees with Wolff that the duct is a trough and not a tube, and proves
it by a satisfactory experiment. He remarks:

[Illustration:

FIG. 87.—Longitudinal section through the head of the honey bee, ♀,
just outside of right antenna: _ant_, antenna with three muscles
attached to _mes_, mesocephalic pillar; _cl_, clypeus; _lbr_,
labrum; 1, chyle-gland (system no. 1, of Siebold); _o_, opening of
the same; _oc_, ocellus; _br_, brain; _n_, neck; _th_, thorax; _oe_,
œsophagus; _s.d_^2, _s.d_^3, common salivary ducts of systems 2 and
3; _v_, salivary valve; _c_, cardo; _ph_, pharynx; _mx′_, labium;
_mx.′p_, labial palpi; _mt_, mentum; _mx_, maxilla; _hyp_,
hypopharynx; _s_, bouton.—After Cheshire.
]

“Bees have the power, by driving blood into the tongue, of forcing
the rod out from the sheath, and distending the wrinkled membrane so
that in section it appears as at _H_, Fig. 86, the membrane assuming
the form of a pouch, given in full length at _A_. It will be seen at
once that this disposition of parts abolishes the side ducts, but
brings the central duct to the external surface. The object of this
curious capability on the part of the bee is, in my opinion, to
permit of cleaning away any pollen grains, or other impediment that
may collect in the side ducts. The membrane is greasy in nature, and
substances or fluids can be removed from it as easily as water from
polished metal. If, now, the sides of a needle, previously dipped
into clove oil in which rosanilin (magenta) has been dissolved, so
as to stain it strongly red, be touched on the centre of the rod,
the oil immediately enters, and passes rapidly upwards and
downwards, filling the trough.”

=Does the hypopharynx represent a distinct segment?=—The facts which
suggest that the hypopharynx may possibly represent a highly
modified pair of appendages, arising from a distinct intermaxillary
segment, are these: Heymons plainly shows that, in the embryo of
Lepisma, the hypopharynx originates as a transverse segment-like
fold in front of the 2d maxillary segment, and larger than it, and
though he does not mention it in his text, it appears like the
rudiment of a distinct segment; the hypopharynx of Ephemeridæ;
arises and remains separate in the nymph from the labium (see
Heymons’ Fig. 29, and there are two lateral projections; see also
Fig. 72, and Vayssiere’s view that it may represent a pair of
appendages; Kolbe also regards it as representing a third pair of
maxillæ, his endolabium, p. 213). Though what is called an unpaired
organ, it is composed of, or supported by, two bilaterally
symmetrical styles, both in Myriopods (Fig. 6, labiella, _stil_) and
in insects (Fig. 77, etc.). On the other hand, in the embryo of
pterygote insects, an intermaxillary segment has not been yet
detected.

LITERATURE OF THE MOUTH-PARTS OR BUCCAL APPENDAGES

_a._ General

=Savigny, Jules-César.= Mémoires sur les animaux sans vertèbres.
1^{re} Part. Description et classification des animaux invertébrés
et articulés, etc. Fasc. 1^{re}. Mém. 1–2. Théorie des organes de la
bouche des crustacés et des insectes. 12 Pl., Paris, 1816, pp.
1–117.

=Gerstfeld, Georg.= Ueber die Mundteile der saugenden Insekten.
Dorpat, 1853.

=Olfers, Ernestus V.= Annotationes ad anatomiam Podurarum. Berolini,
1862, 4 Pls.

=Gerstaecker, Carl Eduard Adolph.= Zur Morphologie der Orthoptera
amphibiotica. (Festschrift zur Feier des hundertjährigen Bestehens
der Gesellschaft naturf. Freunde zu Berlin. 4º, 1873, pp. 39–59, 1
Taf.)

=Muhr, Joseph.= Die Mundteile der Orthoptera. Ein Beitrag zur
vergleichenden Anatomie. (Jahrbuch “Lotos.” Prag, 1877, pp. 40–71, 8
Taf.)

=Burgess, Edward.= The anatomy of the head and the structure of the
maxilla in the Psocidæ. (Proc. Boston Soc. Nat. Hist., xix, 1878,
pp. 291–296, 1 Pl.)

=Meinert, Fr.= Sur la conformation de la tête et sur l’interpretation
des organes buccaux chez les insectes, ainsi que sur la systématique
de cette ordre. (Ent. Tidsskr., 1. Arg., 1880, pp. 147–150.)

—— Tungens udskydelighed hos Steninerne, en slaegt af Staphylinernes
familie. (Vidensk. meddel. fra den naturh. Foren, 1884–1886, pp.
180–207, 2 Pls. Also Zool. Anzeiger, 1887, pp. 136–139.)

=Müller, A.= Vergleichend-anatomische Darstellung der Mundteile der
Insekten. Villach, 1881, 3 Taf.

=Kraepelin, Karl.= Ueber die Mundwerkzeuge der saugenden Insekten.
(Zool. Anzeiger, 1882, pp. 574–579.)

=Dewitz, H.= Ueber die führung an den Körperanhangen der Insekten.
(Berlin. Zeitschr. xxvi., 1882, pp. 51–68, Figs.)

=Wolter, Max.= Die Mundbildung der Orthopteren mit specieller
Berücksichtigung der Ephemeriden. 4 Taf. Greifswald, 1883.

=Oudemans, J. T.= Beiträge zur Kenntniss der Thysanura und Collembola.
(Bijdragen tot de Dierkunde, pp. 149–226. Amsterdam, 1888, 3 Taf.)

=Smith, John B.= An essay on the development of the mouth-parts of
certain insects. (Trans. Amer. Philosophical Soc., xix, pp. 175–198,
3 Pls.)

Also articles by Chatin, McLachlan, Riley, Wood-Mason.

_b._ Thysanoptera (Physapoda)

=Jordan, Karl.= Anatomie und biologie der Physapoda. (Zeitschr. f.
wissens. Zool., xlvii, pp. 541–620, 3 Taf. 1888.)

=Garman, H.= The mouth-parts of the Thysanoptera. (Bull. Essex Inst.,
xxii, 4 pp., Fig. 1890.)

—— The asymmetry of the mouth-parts of Thysanoptera. (Amer. Naturalist,
July, 1896, pp. 591–593, Fig.)

=Bohls, J.= Die Mundwerkzeuge der Physapoden. Dissertation Göttingen,
1891, pp. 1–36.

=Uzel, Heinrich.= Monographie der Ordnung Thysanoptera. Königgrätz,
1895, pp. 472, 10 Taf., 9 Figs.

_c._ Hemiptera

=Léon, N.= Beiträge zur Kenntniss der Mundteile der Hemipteren. Jena,
1887, pp. 47, 1 Taf.

—— Labialtaster bei Hemipteren. (Zool. Anzeiger, pp. 145–147, 1892, 1
Fig.)

—— Beiträge zur Kenntniss des Labiums der Hydrocoren. (Zool. Anzeiger,
März 29, 1897, pp. 73–77, Figs. 1–5.)

=Geise, O.= Mundteile der Rhynchoten. (Archiv f. Naturgesch., xlix,
1883, pp. 315–373, 1 Taf.)

=Wedde, Hermann.= Beiträge zur Kenntniss des Rhynchotenrüssels. (Archiv
f. Naturgesch., li Jahrg., 1 Bd., 1885, pp. 113–148, 2 Taf.)

=Smith, John B.= The structure of the hemipterous mouth. (Science, April
1, 1892, pp. 189–190, Figs. 1–5.)

_d._ Coleoptera

=Smith, John B.= The mouth-parts of _Copris carolina_; with notes on the
homologies of the mandibles. (Trans. Amer. Ent. Soc., xix, April,
1892, pp. 83–87, 2 Pls.)

_e._ Lepidoptera

=Kirbach, P.= Ueber die Mundwerkzeuge der Schmetterlinge. (Zool.
Anzeiger, vi Jahrg., 1883, pp. 553–558, 2 Figs.)

—— Ueber die Mundwerkzeuge der Schmetterlinge. (Archiv f.
Naturgeschichte, 1884, pp. 78–119, 2 Taf.)

=Walter, Alfred.= Palpus maxillaris Lepidopterorum. (Jenaische Zeitschr.
f. Naturwiss, xviii, 1884, pp. 121–173, Taf.)

—— Beiträge zur Morphologie der Lepidoptera. I, Mundteile. (Jenaische
Zeitschr. f. Naturwiss, xviii, 1885, pp. 751–807, 2 Taf.)

=Breitenbach, W.= Vorläufige Mitteilung über einige neue Untersuchungen
an Schmetterlingsrüsseln. (Archiv f. mikroskop. Anatomie, xiv, 1877,
pp. 308–317, 1 Taf.)

—— Untersuchungen an Schmetterlingsrüsseln. (Ibid., xv, 1878, pp. 8–29,
1 Taf.)

—— Ueber Schmetterlingsrüssel. (Entomolog. Nachr. 5 Jahrg., 1879, pp.
237–243, 1 Taf.)

—— Der Schmetterlingsrüssel. (Jenaische Zeitschr. f. Naturwiss, 1881.)

_f._ Siphonaptera

=Kräpelin, K.= Ueber die systematische Stellung der Puliciden.
(Festschrift z. 50 jahr. Jubil. d. Realgymnas. Iohanneum, Hamburg,
pp. 17, 1 Taf. 1884.)

=Kellogg, V. L.= The mouth-parts of the Lepidoptera. (Amer. Nat., xxix,
1895, pp. 546–556, 1 Pl. and Fig.)

_g._ Diptera

=Menzbier, Michael Alexander.= Ueber das Kopfskelett und die
Mundwerkzeuge der Zweiflügler. (Bull. Soc. Imp. Natur. de Moscou,
lv, 1880, pp. 8–71, 2 Taf.)

=Dimmock, George.= The anatomy of the mouth-parts and of the sucking
apparatus of some Diptera. Boston, 1881, pp. 48, 4 Pls.

=Meinert, F.= Fluernes Munddele. Trophi Dipterorum. Kjöbenhavn, 1881, 6
Pls.

—— Die Mundteile der Dipteren. (Zool. Anz. 1882, pp. 570–574, 599–603.)

=Becher, E.= Zur Kenntniss der Mundteile der Dipteren. (Denkschr. Akad.
d. Wissensch. Wien., xlv, 1882, pp. 123–162, 4 Taf.)

=Hansen, H. J.= Fabrica oris dipterorum: Dipterernes mund: anatomisk og
systematisk henseende. 1 Tabanidae, Bombyliidae, Asilidae, Thereva,
Mydas, Apiocera. (Naturhist. Tidsskrift, 1883, xiv, pp. 1–186, Taf.
1–5.)

=Kräpelin, Karl.= Zur Anatomie und Physiologie des Rüssels von Musca.
(Zeitschr. f. wissensch. Zool., xxxix, 1883, pp. 683–719, 2 Taf.)

=McCloskie, George.= Kraepelin’s Proboscis of the house-fly. (American
Naturalist, xviii, 1884, pp. 1234–1244, Figs.)

=Langhoffer, August.= Beiträge zur Kenntniss der Mundtheile der
Dipteren. Jena, 1888, pp. 1–32.

=Smith, John B.= A contribution toward a knowledge of the mouth-parts of
the Diptera. (Trans. Amer. Ent. Soc., xvii, Nov. 1890, pp. 319–339,
Figs. 1–22.)

_h._ Hymenoptera

=Briant, Travers J.= On the anatomy and functions of the tongue of the
honey-bee (worker). (Journ. Linn. Soc., London, xvii, 1884, pp.
408–416, 2 Pls.)

=Breithaupt, P. F.= Ueber die Anatomie und die Funktionen der
Bienenzunge. (Archiv f. Naturgesch., Jahrg. lii, 1886, pp. 47–112, 2
Taf.)

_i._ Larval stages

=Brauer, F.= Die Zweiflügler des kaiserlichen Museums zu Wien. III,
Systematische Studien auf Grundlage der Dipterenlarven nebst einer
Zusammenstellung von Beispielen aus der Litteratur uber dieselben
und Beschreibung neuer Formen. (Denkschr. math.-naturwiss. Cl. k.
Akad. Wiss. Wien, 1883, xlvii, pp. 100, 5 Taf.)

=Dewitz, H.= Ueber die Führung an den Körperanhangen der Insekten
speziell betrachtet an der Legescheide der Acridier, dem Stachel der
Meliponem und den Mundteilen der Larve von Myrmeleon, nebst
Beschreibung dieser Organe. (Berliner ent. Zeitschr., xxvi, 1882,
pp. 51–68.)

—— Die Mundteile der Larve von Myrmeleon. (Sitzungsber. d. Ges.
naturforsch. Freunde zu Berlin, 1881, pp. 163–166.)

=Redtenbacher, Josef.= Uebersicht der Myrmeleonidenlarven. (Denkschrift,
math.-naturwiss. Cl. k. Akad. Wiss. Wien, 1884, xlviii, pp. 335–368,
7 Taf.)

=Schiödte, J. G.= De metamorphosi Eleutheratorum. Bidrag til insekternes
udviklingshistorie. (Kroyer’s Naturhist. Tidsskrift. Kjöbenhavn. 12
Teile mit 88 Taf., 1862–1883.)

_j._ Embryonic stages

=Heymons, Richard.= Grundzüge der Entwicklung und des Körpersbaues von
Odonaten und Ephemeridem (Anhang zu den Abhandl. K. Akad. d.
Wissens. Berlin, 1896, p. 22, 2 Taf. See Figs. 5, 29.)

—— Entwicklungsgeschichtliche Untersuchungen an _Lepisma saccharina_ L.
(Zeitschr. f. Wissens. Zoologie, lxii, 1897, p. 595, 2 Taf. See Fig.
10.)

THE THORAX AND ITS APPENDAGES

_a._ The thorax; its external anatomy

The middle region of the body is called the thorax, and in general
consists of three segments, which are respectively named the
_prothorax_, _mesothorax_, and _metathorax_ (Figs. 88, 89, 98).

[Illustration:

FIG. 88.—External anatomy of _Melanoplus spretus_, the head and thorax
disjointed.
]

The thorax contains the muscles of flight and those of the legs, besides
the fore intestine (œsophagus and proventriculus), as well as, in the
winged insects, the salivary glands.

In the more generalized orders, notably the Orthoptera, the three
segments are distinct and readily identified.

[Illustration:

FIG. 89.—Locust, Melanoplus, side view, with the thorax separated from
the head and abdomen, and divided into its three segments.
]

Each segment consists of the _tergum_, _pleurum_, and _sternum_. In the
prothorax these pieces are not subdivided, except the pleural; in such
case the tergum is called the _pronotum_. The prothorax is very large in
the Orthoptera and other generalized forms, as also in the Coleoptera,
but small and reduced in the Diptera and Hymenoptera. In the winged
forms the tergum of the mesothorax is differentiated into four pieces or
plates (sclerites). These pieces were named by Audouin, passing from
before backwards, the _præscutum_, _scutum_, _scutellum_, and
_postscutellum_. In the nymph stage and in the wingless adults of
insects such as the Mallophaga, the true lice, the wingless Diptera,
ants, etc., these parts by disuse and loss of the wings are not
differentiated. It is therefore apparent that their development depends
on that of the muscles of flight, of which they form the base of
attachment. The scutum is invariably present, as is the scutellum. The
former in nearly all insects constitutes the larger part of the tergum,
while the latter is, as its name implies, the small shield-shaped piece
directly behind the scutum.

[Illustration:

FIG. 90.—Thorax of _Telea polyphemus_, side view, pronotum not
represented: _em_, epimerum of prothorax, the narrow piece above
being the prothoracic episternum; _ms_, mesoscutum; _scm_,
mesoscutellum; _ms″_, metascutum; _scm‴_, metascutellum; _pt_, a
supplementary piece near the insertion of tegulæ; _w_, pieces
situated at the insertion of the wings, and surrounded by membrane;
_epm″_, episternum of the mesothorax; _em″_, epimerum of the same;
_epm‴_, episternum of the metathorax; _em‴_ epimerum of the same,
divided into two pieces; _c′_, _c″_, _c‴_, coxæ; _te′_, _te″_,
_te‴_, trochantines; _tr_, _tr_, _tr_, trochanters. _A_, tergal view
of the mesothorax of the same; _prm_, præscutum; _ms_, scutum;
_scm_, scutellum; _ptm_, postscutellum; _t_, tegula.
]

The præscutum and postscutellum are usually minute and crowded down out
of sight between the opposing segments. As seen in Fig. 90, the
præscutum of most moths (Telea) is a small rounded piece, bent
vertically down so as not to be seen from above. In Polystœchotes and
also in Hepialus the præscutum is large, well-developed, triangular, and
wedged in between the two halves of the scutum. The postscutellum is
still smaller, usually forming a transverse ridge, and is rarely used in
taxonomy.

[Illustration:

FIG. 91.—Thorax of the house-fly: _prn_, pronotum; _prsc_, præscutum;
_sc′_, mesoscutum; _sct′_, mesoscutellum; _psct′_, postscutellum;
_al_, insertion of squama, extending to the insertion of the wings,
which have been removed; _msphr_, mesophragma; _h_, balancer
(halter); _pt_, tegula; _mtn_, metanotum; _epis_, _epis′_, _epis″_,
episternum of pro-, meso-, and metathorax; _epm′_, _epm″_, meso- and
meta-epimerum; _st′_, _st″_, meso- and metasternum; _cx′_, _cx″_,
_cx‴_, coxæ; _tr′_, _tr″_, _tr‴_, trochanters of the three pairs of
legs; _sp′_, _sp″_, _sp‴_, _sp‴′_, _sp‴″_, first to fifth spiracles;
_tg′_, _tg″_, tergites of first and second abdominal segments; _u′_,
_u″_, urites.
]

The metathorax is usually smaller and shorter than the mesothorax, being
proportioned to the size of the wings. In certain Neuroptera and in
Hepialidæ and some tineoid moths, where the hind wings are nearly as
large as those of the anterior pair, the metathorax is more than half or
nearly two-thirds as large as the mesothorax. In Hepialidæ the præscutum
is large and distinct, while the scutum is divided into two widely
separated pieces. The postscutellum is nearly or quite obsolete.

The pleurum in each of the three thoracic segments is divided into two
pieces; the one in front is called the _episternum_, since it rests upon
the sternum; the other is the _epimerum_. To these pieces, with the
sternum in part, the legs are articulated (Fig. 89).

Between the episterna is situated the breastplate or _sternum_, which is
very large in the more primitive forms, as the Orthoptera, and is small
in the Diptera and Hymenoptera.

[Illustration:

FIG. 92.—Prothorax of _Geometra papilionaria_: _n_, notum; _p_,
pleura; _st_, sternum; _pt_, patagia; _m_, membrane; _f_, femur;
_h_, a hook bent backwards and beneath, and connecting the pro- with
the mesothorax.—After Cholodkowsky.
]

The episterna and epimera are in certain groups, Neuroptera, etc.,
further subdivided each into two pieces (Fig. 102). The smaller pieces,
hinging upon each other and forming the attachments of the muscles of
flight, differ much in shape and size in insects of different orders.
The difference in shape and degree of differentiation of these parts of
the thorax is mentioned and illustrated under each order, and reference
to the figures will obviate pages of tedious description. A glance,
however, at the thorax of a moth, fly, or bee, where these numerous
pieces are agglutinated into a globular mass, will show that the
spherical shape of the thorax in these insects is due to the enlargement
of one part at the expense of another; the prothoracic and metathoracic
segments being more or less atrophied, while the mesothorax is greatly
enlarged to support the powerful muscles of flight, the fore wings being
much larger than those appended to the metathorax. In the Diptera, whose
hinder pair of wings are reduced to the condition of halteres, the
reduction of the metathorax as well as prothorax is especially marked
(Fig. 91).

=The patagia.=—On each side of the pronotum of Lepidoptera are two
transversely oval, movable, concavo-convex, erectile plates, called
_patagia_ (Fig. 92). On cutting those of a dry Catocala in two, they
will be seen to be hollow. Cholodkowsky[19] states that they are filled
with blood and tracheal branches; and he went so far as to regard them
as rudimentary prothoracic wings, in which view he was corrected by
Haase,[20] who compares them with the tegulæ, regarding them also as
secondary or accessory structures.

=The tegulæ.=—On the mesothorax are the _tegulæ_ of Kirby (_pterygodes_
of Latreille, _paraptera_ of McLeay, _hypoptère_ or _squamule_), which
cover the base of the fore wings, and are especially developed in the
Lepidoptera (Fig. 90, _A_, _t_) and in certain Hymenoptera (Fig. 95,
_c_).

The external opening of the spiracles just under the fore wings, is
situated in a little plate called by Audouin the _peritreme_.

[Illustration:

FIG. 93.—Transformation of the bumble bee, Bombus, showing the
transfer of the 1st abdominal larval segment (_c_) to the thorax,
forming the propodeum of the pupa (_D_) and imago; _n_, spiracle of
the propodeum. _A_, larva; _a_, head; _b_, 1st thoracic; _c_, 1st
abdominal segment. _B_, semipupa; _g_, antenna; _h_, maxillæ; _i_,
1st; _j_, 2d leg; _k_, mesoscutum; _l_, mesoscutellum; _m_,
metathorax; _d_, urite (sternite of abdomen); _e_, pleurite; _f_,
tergite; _o_, ovipositor; _r_, lingua; _q_, maxilla.
]

In the higher or aculeate Hymenoptera, besides the three segments
normally composing the thorax, the basal abdominal segment is during the
change from the larva to the pupa transferred to this region, making
four segments. This first abdominal is called “the median segment”
(Figs. 93–95). In such a case the term _alitrunk_ has been applied to
this region, _i.e._ the thorax, as thus constituted. Latreille wrongly
stated that in the Diptera the first abdominal segment also entered into
the composition of the thorax; but Brauer has fully disproved that view,
as may be seen by an examination of his sketches which we have copied
(Fig. 94).

[Illustration:

FIG. 94.—7, 8, thorax of _Tipula gigantea_; 9, of Leptis; 10, thorax
of _Tabanus bromius_ after the removal of the abdomen, in order to
bring into view the inner mesophragma (_f_), and to show the
extension of the metathorax _g_ and _g′_; _tr_, trochanter; 11, hind
end of the mesothorax, the entire metathorax, and the 1st and 2d
abdominal segments of _Volucella zonaria_, seen from the side. The
internal mesophragma (_f_), and the position of the muscle inserted
in it, are indicated by the two lines _M_. _p_, Callus postalaris;
_pr_ (_pz_ in 8), callus præalaris Osten Sacken (= “patagium” of
some authors); _g_, metanotum; _g′_, metepimerum, “segment médiaire”
of Latreille (wrongly considered by him to be the 1st abdominal
segment); 4, metasternum (hypopleura of Osten Sacken); 5 (?
“episternum of metathorax” (Brauer) = metapleura of Osten Sacken);
6, and also _H_, halter; _st_^1, mesothoracic stigma; _st_^2,
metathoracic stigma; _st_^3, first abdominal stigma; γ,
dorsopleural; δ, sternopleural; ε, mesopleural sutures; _h_, 1st,
_i_, 2d, abdominal segment; _al_, wing; _alul_, alula. 12, the head
and the three thoracic rings, and the 1st abdominal segment of
_Ephemera vulgata_, the connecting membranes are in white: _a_,
prothorax; _b_, præscutum; _c_, scutum; _d_, scutellum; _e_,
postscutellum; _ps_, postscutellum of mesothorax.—After Brauer.
]

[Illustration:

FIG. 95.—Alitrunk of _Sphex chrysis_: _A_, dorsal aspect; _a_,
pronotum; _b_, mesonotum; _c_, tegula; _d_, base of fore,—_e_, of
hind, wing; _f_, _g_, divisions of metanotum; _h_, median (true
first abdominal) segment; _i_, its spiracle; _k_, second abdominal
segment, usually called the petiole or first abdominal segment. _B_,
posterior aspect of the median segment; _a_, upper part; _b_,
superior,—_c_, inferior, abdominal foramen; _d_, ventral plate of
median segment; _e_, coxa.—After Sharp.
]

The sternum is in rare cases subdivided into two halves, as in the meso-
and metathorax of the cockroach; in Forficula the prosternum is divided
into four pieces besides the sternum proper (Fig. 96); and in Embia,
also, the sternites, according to Sharp, are complex.

[Illustration:

FIG. 96.—Sternal view of pro-, meso-, and metathorax of _Forficula
tæniata_: _pst_, præsternum, divided into 4 pieces; _st_, pro-,
_st′_, meso-, _st″_, metasternum; _cx_, coxa; _not_, notum.
]

[Illustration:

FIG. 97.—_A_, under surface of prothorax, or prosternum, of _Dyticus
circumflexis_: 2._g_, prosternum; 2._f_, episternum; 2._h_,
epimerum; 2._s_, antefurca or entothorax.
]

[Illustration:

FIG. 98.—Meso- (_G__{2}) and metathoracic ganglia (_G__{1}), with the
apodemes of Gryllotalpa.—After Graber.
]

[Illustration:

FIG. 99.—Parts of the mesothorax of Dyticus: _A_, mesosternum; 3._a_,
præscutum; 3._b_, scutum; 3._c_, scutellum; 3._d_, postscutellum;
3._e_, parapteron; 3._g_, mesosternum; 3._f_, episternum; 3._h_,
epimerum; 3._s_, medifurca or entothorax.
]

[Illustration:

FIG. 100.—Parts of the metathorax of Dyticus: _A_, metasternum; 4._a_,
præscutum; 4._b_, scutum; 4._c_, scutellum; 4._d_, postscutellum;
4._e_, parapteron; 4._f_, episternum; 4._g_, metasternum; 4._h_,
epimerum; 4._s_, postfurca.—This and Figs. 97 and 99 from Audouin,
after Newport.
]

=The apodemes.=—The thorax is supported within by beam-like processes,
or _apodemes_, which pass inward and also form attachments for the
muscles. Those passing up from the sternum form the _entothorax_ of
Audouin, and the process of each thoracic segment is called respectively
the _antefurca_, _medifurca_, and _postfurca_. In the Orthoptera
(Caloptenus and Anabrus), the antefurca is large, thin, flattened,
directed forward, and bounds each side of the prothoracic ganglion. In
the Coleoptera two plates (Fig. 97, 2._s_) arise from the inside of the
sternum and “form a collar or leave a circular hole between them for the
passage of the nervous cord” (Newport). The medifurca is a pair of flat
processes which diverge and bridge the commissure, while the postfurca
is situated under the commissure. In beetles (Dyticus) Newport states
that it is expanded into two broad plates, to which the muscles of the
posterior legs are attached. Graber also notices in the mole cricket
between the apodemes of the meso- and metathorax, a flattened spine
(Fig. 98, _do_) with two perforations through which pass the commissures
connecting the ganglia. Besides these processes there are large, thin,
longitudinal partitions passing down from the tergum (or dorsum), called
_phragmas_; they are most developed in those insects which fly best,
_i.e._ in Coleoptera (Figs. 97–101), Lepidoptera, Diptera, and
Hymenoptera, none being developed in the prothorax. (The term _phragma_
has also been applied to a partition formed by the inflexed hinder edge
of this segment, and is present only in those insects in which the
prothorax is movable.—Century Dictionary.) All these ingrowths may be in
general termed _apodemes_. There are similar structures in Crustacea and
also in Limulus; but Sharp restricts this term to minute projections in
beetles (Goliathus) situated at the sides of the thorax near the wings.
(Insecta, p. 103, Fig. 57.) The internal processes arising from the
sternal region have been called _endosternites_.

[Illustration:

FIG. 101.—Internal skeleton of _Lucanus cervus_, ♂, head: _A_,
antenna; _f_, mandible; _d_, mentum; 2, 4, tendons of mandible; _f_,
_u_, _t_, parts of the tentorium; 3 _e_, labial muscles. Thorax: 2,
prothorax; 3, 4, meso- and metathorax fused solidly together; 3 _r_,
acetabulum of prothorax, into which the coxa is inserted; 2 _s_,
sternum; 3_t_, acetabulum of mesothorax, 4_r_, of metathorax; 3 _s_,
mesothoracic sternum fused with that of the metathorax (4_g_); 4
_s_, apodeme.—After Newport.
]

=The acetabula.=—These are the cavities in which the legs are inserted.
They are situated on each side of the posterior part of the sternum, in
each of the thoracic segments. They are, in general, formed by an
approximation of the sternum and epimerum, and sometimes, also, of the
episternum, as in Dyticus (Fig. 97, _A_). This consolidation of parts,
says Newport, gives an amazing increase of strength to the segments, and
is one of the circumstances which enables the insect to exert an
astonishing degree of muscular power.

TABULAR VIEW OF THE SEGMENTS, PIECES, AND APPENDAGES OF THE THORAX

═══════════════════════╤═══════════════════════╤═══════════════════════
NAME OF SEGMENT │ PIECES (SCLERITES) │ APPENDAGES
───────────────────────┼───────────────────────┼───────────────────────
1. Prothorax │Pronotum, sometimes │
│ differentiated into │
│Scutum │1st pair of legs
│Scutellum │Patagia
│Episternum │
│Epimerum │
│Sternum │
│Antefurca │
│ │
2. Mesothorax │Præscutum │
│Scutum │2d pair of legs
│Scutellum │1st pair of wings
│Proscutellum │Tegulæ
│Episternum │Squamæ (Alulæ)
│Epimerum │Peritreme
│Sternum │
│Mesofurca │
│Mesophragma │
│Apodemes │
│ │
3. Metathorax │Præscutum │
│Scutum │3d pair of legs
│Scutellum │2d pair of wings
│Postscutellum │(Halteres of Diptera)
│Episternum │
│Epimerum │
│Sternum │
│Postfurca │
│Metaphragma │
│Apodemes │
───────────────────────┴───────────────────────┴───────────────────────

[Illustration:

FIG. 102.—External anatomy of the trunk of _Hydröus piceus_: _A_,
sternal—_B_, tergal aspect; 2, pronotum; 2 _a_, prosternum; 2 _f_,
episternum; 3 _a_, præscutum; 3 _b_, scutum; 3 _c_, scutellum; 3
_d_, postscutellum; 3 _g_, mesosternum; 3 _h_, episternum; 3 _f_,
epimerum; 3 _i_, crest of the mesosternum; 3 _a_, parapteron; 3 _k_,
coxa; 4 _a_, metapræscutum; 4 _b_, metascutum; 4 _c_, metascutellum;
4 _d_, postscutellum; 4 _e_, tegula; 4 _f_, episternum; 4 _h_,
epimerum; 4 _g_, metasternum; 4 _i_, crest of metasternum; 4 _k_ and
_l_, coxa; 4 _m_, trochanter; _n_, femur; _o_, tibia; _p_, tarsus;
_q_, unguis; 7–11, abdominal segments.—After Newport.
]

_b._ The legs: their structure and functions

The mode of insertion of the legs to the thorax is seen in Figs. 90, 97,
101, and 103. They are articulated to the episternum, epimerum, and
sternum, taken together, and consist of five segments. The basal segment
or joint is the _coxa_, situated between the episternum and trochanter.
The coxa usually has a posterior subdivision or projection, the
_trochantine_; sometimes, as in Mantispa (Fig. 103), the trochantine is
obsolete. We had previously supposed that the trochantine was a separate
joint, but now doubt whether it represents a distinct segment of the
leg, and regard it as only a subdivision of the coxa. It is attached to
the epimerum, and is best developed in Panorpidæ, Trichoptera, and
Lepidoptera. In the Thysanura the trochantine is wanting, and in the
cockroach it merely forms a subdivision of the coxa, its use being to
support the latter. The second segment is the trochanter, a more or less
short spherical joint on which the leg proper turns; in the parasitic
groups (Ichneumonidæ, etc., Fig. 104) it is usually divided into two
pieces, though there are some exceptions. The trochanter is succeeded by
the _femur_, _tibia_, and _tarsus_, the latter consisting of from one to
five segments, the normal number being five. Tuffen West believed that
the pulvillus is the homologue of an additional tarsal joint, “a sixth
tarsal joint.” The last tarsal segment ends in a pair of freely movable
claws (ungues), which are modified setæ; between the claws is a
cushion-like pad or adhesive lobe, called the _empodium_ or _pulvillus_
(Fig. 105, also variously called _arolium_, _palmula_, _plantula_,
_onychium_, its appendage being called _paronychium_ and also
_pseudonychium_). It is cleft or bilobate in many flies, but in Sargus
trilobate. All these parts vary greatly in shape and relative size in
insects of different groups, especially Trichoptera, Lepidoptera,
Diptera, and Hymenoptera. In certain flies (_e.g._ Leptogaster) the
empodium is wanting (Kolbe). By some writers the middle lobe is called
the empodium and the two others pulvilli.

[Illustration:

FIG. 103.—Side view of meso- and metathorax of _Mantispa brunnea_,
showing the upper and lower divisions of the epimerum (_s. em′_, _s.
em″_, _i. em′_, _i. em″_); _s. epis_, _i. epis″_, the same of the
episternum.
]

[Illustration:

FIG. 104.—Divided (ditrochous) trochanter of an ichneumon: _cx_, coxa;
_tr_, the two divisions of the trochanter; _f_, femur.—After Sharp.
]

The fore legs are usually directed forward to drag the body along, while
the middle and hind legs are directed outward and backward to push the
body onwards. While arachnids walk on the tip ends of their feet,
myriopods, Thysanura, and all larval insects walk on the ends of the
claws, but insects generally, especially the adults, are, so to speak,
plantigrade, since they walk on all the tarsal joints. In the aquatic
forms the middle and hind tarsi are more or less flattened, oar-like,
and edged with setæ. In leaping insects, as the locusts and
grasshoppers, and certain chrysomelids, the hind femora are greatly
swollen owing to the development of the muscles within. The tibia,
besides bearing large, lateral, external spines, occasionally bears at
the end one or more spines or spurs called _calcaria_. The fore tibia
also in ants, etc., bear tactile hairs, and chordotonal organs, as well
as other isolated sense-organs (Janet), and, in grasshoppers, ears.

In the Carabidæ the legs are provided with combs for cleaning the
antennæ (Fig. 107), and in the bees and ants these cleansing organs are
more specialized, the pectinated spine (_calcar_) being opposed by a
tarsal comb (Fig. 106, _d_; for the wax-pincers of bees, see _g_). In
general the insects use their more or less spiny legs for cleansing the
head, antennæ, palpi, wings, etc., and the adaptations for that end are
the bristles or spinules on the legs, especially the tibiæ.

[Illustration:

FIG. 105.—Foot of honey-bee, with the pulvillus in use: _A_, under
view of foot; _t_, _t_, 3d–5th tarsal joints; _a n_, unguis; _f h_,
tactile hairs; _p v_, pulvillus; _cr_, curved rod. _B_, side view of
foot. _C_, central part of sole; _pd_, pad; _cr_, curved rod; _pv_,
pulvillus unopened.—After Cheshire.
]

[Illustration:

FIG. 106.—Modifications of the legs of different bees. _A_, Apis: _a_,
wax-pincer and outer view of hind leg; _b_, inner aspect of
wax-pincer and leg, with the nine pollen-brushes or rows of hairs;
_c_, compound hairs holding grains of pollen; _d_, anterior leg,
showing antenna-cleaner; _e_, spur on tibia of middle leg. _B_,
Melipona: _f_, peculiar group of spines at apex of tibia of hind
leg; _g_, inner aspect of wax-pincer and first tarsal joint. _C_,
Bombus: _h_, wax-pincer; _i_, inner view of the same and first
tarsal joint, all enlarged.—From _Insect Life_, U. S. Div. Ent.
]

Osten Sacken states that among Diptera the aerial forms (Bombylidæ,
etc.) with their large eyes or holoptic heads, which carry with them the
power of hovering or poising, have weak legs, principally fit for
alighting. On the other hand, the pedestrian or walking Diptera
(Asilidæ, etc.) “use the legs not for alighting only, but for running,
and all kinds of other work, seizing their prey, carrying it, climbing,
digging, etc.; their legs are provided not only with spines and
bristles, but with still other appendages, which may be useful, or only
ornamental, as secondary sexual characters.”

[Illustration:

FIG. 107.—End of tibia and tarsal joints of Anophthalmus; _c_, comb.
]

=Tenent hairs.=—Projecting from the lower surface of the empodium are
the numerous “tenent hairs,” or holding hairs, which are modified
glandular setæ swollen at the end and which give out a minute quantity
of a clear adhesive fluid (Figs. 108, 109, 130, 134). In larval insects,
and the adults of certain beetles, Coccidæ, Aphidæ, and Collembola,
which have no empodium, there are one or more of these tenent hairs
present. They enable the insect to adhere to smooth surfaces.

[Illustration:

FIG. 108.—Transverse section through a tarsal joint of Telephorus, a
beetle: _ch_, cuticula of the upper side; _m_, its matrix; _ch′_,
the sole; _m′_, its matrix; _h_, adhesive hair; _h′_, tactile hair,
supplied with a nerve (_n′_), and arising from a main nerve (_n_);
_n″_, ganglion of a tactile hair; _t_, section of main trachea, from
which arises a branch (_t′_); _dr_, glands which open into the
adhesive hairs, and form the sticky secretion; _e_, chitinous
thickening; _s_, sinew; _b_, membrane dividing the hollow space of
the tarsal joint into compartments. See p. 111.—After Dewitz.
]

Striking sexual secondary characters appear in the fore legs of the male
Hydrophilus, the insect, as Tuffen West observes, walking on the end of
the tibia alone and dragging the tarsus after it. The last tarsal joint
is enlarged into the form of an irregular hollow shield. The most
completely suctorial feet of insects are those of the anterior pair of
Dyticus (Fig. 132). The under side of the three basal joints is fused
together and enlarged into a single broad and nearly circular shield,
which is convex above and fringed with fine branching hairs, and covered
beneath with suckers, of which two are exceptionally large; by this
apparatus of suckers the male is enabled to adhere to the back of its
mate during copulation. The line branching hairs around the edge prevent
the water from penetrating and thus destroying the vacuum, “while if the
female struggle out of the water, by retaining the fluid for some time
around the sucker, they will in like manner under these altered
conditions equally tend to preserve the effectual contact.” (Tuffen
West.)

[Illustration:

FIG. 109.—Cross-section through tarsus of a locust: _ch_, cuticula of
upper side,—_ch′_, _ch″_, _ch‴_, of sole; _ch_, tubulated layer;
_ch″_, lamellate layer; _ch‴_, inner projections of _ch″_. Other
lettering as in Fig. 101. See p. 113.—After Dewitz.
]

In the saw-flies (Uroceridæ and Tenthredinidæ) and other insects, there
are small membranous oval cushions (_arolia_, Figs. 109 and 131) beneath
each or nearly each tarsal joint.

The triunguline larvæ of the Meloidæ are so called from apparently
having three ungues, but in reality there is only a single claw,
with a claw-like bristle on each side.

=Why do insects have but six legs?=—Embryology shows that the
ancestors of insects were polypodous, and the question arises to
what cause is due the process of elimination of legs in the
ancestors of existing insects, so that at present there are no
functional legs on the abdomen, these being invariably restricted
(except in caterpillars) to the thorax, and the number never being
more than six. It is evident that the number of six legs was fixed
by heredity in the Thysanura, before the appearance of winged
insects. We had thought that this restriction of legs to the thorax
was in part due to the fact that this is the centre of gravity, and
also because abdominal legs are not necessary in locomotion, since
the fore legs are used in dragging the insect forwards, while the
two hinder pairs support and push the body on. Synchronously with
this elimination by disuse of the abdominal legs, the body became
shortened, and subdivided into three regions. On the other hand, as
in caterpillars, with their long bodies, the abdominal legs of the
embryo persist; or if it be granted that the prop-legs are secondary
structures, then they were developed in larval life to prop up and
move the abdominal region.

The constancy of the number of six legs is explained by Dahl as
being in relation to their function as climbing organs. One leg, he
says, will almost always be perpendicular to the plane when the
animal is moving up a vertical surface; and, on the other hand, we
know that three is the smallest number with which stable equilibrium
is possible; an insect must therefore have twice this number, and
the great numerical superiority of the class may be associated with
this mechanical advantage. (This numerical superiority of insects,
however, seems to us to be rather due to the acquisition of wings,
as we have already stated on pages 2 and 120.)

=Loss of limbs by disuse.=—Not only are one or both claws of a single
pair, or those of all the feet atrophied by disuse, but this process of
reduction may extend to the entire limb.

In a few insects one of the claws of each foot is atrophied, as in
the feet of the Pediculidæ, of many Mallophaga, all of the Coccidæ,
in Bittacus, Hybusa (Orthoptera), several beetles of the family
Pselaphidæ, and a weevil (Brachybamus). Hoplia, etc., bear but a
single claw on the hind feet, while the allied Gymnoloma has only a
single claw on all the feet. Cybister has in general a single
immovable claw on the hind feet, but _Cybister scutellaris_ has,
according to Sharp, on the same feet an outer small and movable
claw. In the water bugs, Belostoma, etc., the fore feet end in a
single claw, while in others (Corisa) both claws are wanting on the
fore feet. Corisa also has no claws on the hind feet; Notonecta has
two claws on the anterior four feet, but none on the hind pair. In
Diplonychus, however, there are two small claws present. (Kolbe.)

[Illustration:

FIG. 110.—Last tarsal joint of _Melolontha vulgaris_, drawn as if
transparent to show the inner mechanism: _un_, claws; _str_,
extensor plate; _s_, tendon of the flexor muscle; _vb_, elastic
membrane between the extensor plate and the sliding surface _u_;
_krh_, process of the ungual joint; _emp_, extensor spine, and _th_,
its two tactile hairs.—After Ockler, from Kolbe.
]

Among the Scarabæidæ, the individuals of both sexes of the fossorial
genus Ateuchus (_A. sacer_) and eight other genera, among them
_Deltochilum gibbosum_ of the United States, have no tarsi on the
anterior feet in either sex. The American genera Phanæus (Fig. 111),
Gromphas, and Streblopus have no tarsal joints in the male, but they are
present in the female, though much reduced in size, and also wanting,
Kolbe states, in many species of Phanæus. The peculiar genus
Stenosternus not only lacks the anterior feet, but also those of the
second and third pair of legs are each reduced to a vestige in the shape
of a simple, spur-like, clawless joint. The ungual joint is wanting in
the weevil Anoplus, and becomes small and not easily seen in four other
genera.

Ryder states that the evidence that the absence of fore tarsi in
Ateuchus is due to the inheritance of their loss by mutilation is
uncertain. Dr. Horn suggests that cases like Ateuchus and
Deltochilum, etc., “might be used as an evidence of the persistence
of a character gradually acquired through repeated mutilation, that
is, a loss of the tarsus by the digging which these insects
perform.” On the other hand, the numerous species of Phanæus do
quite as much digging, and the anterior tarsi of the male only are
wanting. “It is true,” he adds, “that many females are seen which
have lost their anterior tarsi by digging; have, in fact, worn them
off; but in recently developed specimens the front tarsi are always
absent in the males and present in the females. If repeated
mutilation has resulted in the entire disappearance of the tarsi in
one fossorial insect, it is reasonable to infer that the same
results should follow in a related insect in both sexes, if at all,
and not in the male only. It is evident that some other cause than
inherited mutilation must be sought for to explain the loss of the
tarsi in these insects.” (Proc. Amer. Phil. Soc., Philadelphia,
1889, pp. 529, 542.)

[Illustration:

FIG. 111.—Fore tibia of _Phanæus carnifex_, ♂, showing no trace of the
tarsus.
]

[Illustration:

FIG. 112.—Fore leg of the mole-cricket: _A_, outer, _B_, inner,
aspect; _e_, ear-slit.—After Sharp.
]

The loss of tarsi may be due to disuse rather than to the inheritance of
mutilations. Judging by the enlarged fore tibiæ, which seem admirably
adapted for digging, it would appear as if tarsi, even more or less
reduced, would be in the way, and thus would be useless to the beetles
in digging. Careful observations on the habits of these beetles might
throw light on this point. It may be added that the fore tarsi in the
more fossorial Carabidæ, such as Clivina and Scarites, as well as those
of the larva of Cicada and those of the mole crickets (Fig. 112), are
more or less reduced; there is a hypertrophy of the tibiæ and their
spines. The shape of the tibia in these insects, which are flattened
with several broad triangular spines, bears a strong resemblance to the
nails or claws of the fossorial limbs of those mammals which dig in hard
soil, such as the armadillo, manis, aardvark, and Echidna. The principle
of modification by disuse is well illustrated in the following cases.

In many butterflies the fore legs are small and shortened, and of little
use, and held pressed against the breast. In the Lycænidæ the fore tarsi
are without claws; in Erycinidæ and Libytheidæ the fore legs of the
males are shortened, but completely developed in the females, while in
the Nymphalidæ the fore legs in both sexes are shortened, consisting in
the males of one or two joints, the claws being absent in the females.
Among moths loss of the fore tarsi is less frequent. J. B. Smith[21]
notices the lack of the fore tarsi in the male of a deltoid, _Litognatha
nubilifasciata_ (Fig. 113), while the hind feet of _Hepialus hectus_ are
shortened. In an aphid (_Mastopoda pteridis_, Esl.) all the tarsi are
reduced to a single vestigial joint (Fig. 114).

[Illustration:

FIG. 113.—Leg of Litognatha: _cx_, coxa; _f_, femur; _t_, tibia; _ep_,
its epiphysis, and _sh_, its shield-like process. The tarsus
entirely wanting.—After Smith.
]

Entirely legless adult insects are rare, and the loss is clearly seen to
be an adaptation due to disuse; such are the females of the Psychidæ,
the females of several genera of Coccidæ (Mytilaspis, etc.), and the
females of the Stylopidæ.

Apodous larval insects are common, and the loss of legs is plainly seen
to be a secondary adaptive feature, since there are annectant forms with
one or two pairs of thoracic legs. All dipterous and siphonapterous
larvæ, those of all the Hymenoptera except the saw-flies, a few
lepidopterous larvæ, some coleopterous, as those of the Rhyncophora,
Buprestidæ, Eucnemidæ, and other families, and many Cerambycidæ are
without any legs. In _Eupsalis minuta_, belonging to the Brenthidæ, the
thoracic legs are minute.

The legs of larvæ end in a single claw, upon the tips of which the
insect stands in walking.

_c._ Locomotion (walking, climbing, and swimming)

=Mechanics of walking.=—To Graber we owe the best exposition of the
mechanics of walking in insects.

“The first segment of the insect leg,” he says, “upon which the
weight of the body rests first of all, is the coxa. Its method of
articulation is very different from that of the other joints. The
enarthrosis affords the most extensive play, particularly in the
Hymenoptera and Diptera.”

In the former the development of their social conditions is very
closely connected with the freest possible use of the legs, which
serve as hands. In the beetles, however, which are very compactly
built, there exists a solid articulation whereby the entire hip
rests in a tent-like excavation of the thorax, and can only be
turned round a single axis, as may be seen in Fig. 115, where _c_
represents the imaginary revolving axis and _d_ the coxa. In the
case we are supposing, therefore, only a backward and forward
movement of the coxa is possible, the extent of the play of which
depends on the size of the coxal pan, as well as certain groin or
bar-like structures which limit further rotation. In the very
dissimilar arrangement which draws in the fore, middle, and hind
legs toward the body it is self-evident that their extent of action
is also different. This arrangement seems to be most yielding on the
fore legs, where the hips, to confine ourselves to the stag-beetles,
can be turned backward and forward 60° from the middle or normal
position, and therefore describe on the whole a curve of 120°. The
angle of turning on the middle leg hardly exceeds a legitimate
limit, yet a forward as well as a backward rotation takes place. The
former is entirely wanting in the hind hips; they can only be moved
backward.

[Illustration:

FIG. 114.—Leg of an Aphid, with the tarsus (_t_) much reduced: 1, 2,
3, legs of 1st, 2d, and 3d pairs.
]

The number and strength of the muscles on which the rotation of the
hips depends, correspond with these varying movements of the
individual legs. Thus, according to Straus Durckheim, the fore coxa
of many beetles possesses five separate muscles and four forward and
one backward roll; the middle coxa a like number of muscles but only
two forward rolls, while the hind hips succeed in accomplishing each
of the motions named with a single muscle.

One can best see how these muscles undertake their work, and above
all how they are situated, if he lays bare the prothorax of the stag
beetle (Fig. 116). Here may be seen first the thick muscle which
turns to the front the rotating axis in its cylindrical pan, and
thus helps to extend the leg, while two other tendons, which take
the opposite direction, are fitted for reflex movements.

[Illustration:

FIG. 115.—Mechanics of an insect’s leg: _d_, coxa,—_c_, axis of
revolution; _a_ and _b_, the coxal muscles; _e_, trochanter muscle
(elevator of the femur); _f_, extensor,—_g_, flexor, of the tibia
(_pn_); _n_, tibial spine; _h_, flexor.—_i_, extensor, of the
foot; _k_, extensor,—_l_, flexor, of the claw; _po_, place of
flexure of the tibia; _p^1q_, leg after being turned back by the
coxa.—_p^1r_, by the simultaneous flexure of the tibia. The
resulting motion of the end of the tibia, through the simultaneous
movement (_no_) and revolution (_nq_), indicates the curve
_nr_.—After Graber.
]

In Fig. 115 the muscles mentioned above, and their modes of working,
may be distinguished by the arrows _a_ and _b_.

In order to simplify matters, we will imagine the second component
part of the normal insect leg, _i.e._ the trochanter (Figs. 116,
117, _r_), as grown together with the third lever, _i.e._ the femur,
as the movement of both parts mostly takes place uniformly.

[Illustration:

FIG. 116.—Section of the fore leg of a stag-beetle, showing the
muscles: _S_, extensor,—_B_, flexor, of the leg; _s_,
extensor,—_b_, flexor, of the femur; _o_, femur; _u_, tibia; _f_,
tarsus; _k_, claw; 109, _s_, extensor,—_b_, flexor, of the
femoro-tibial joint, both enlarged.—After Graber.
]

The pulling of the small trochanter muscle works against the weight
of the body when this is carried over on to the trochanter by means
of the coxa, as seen at the arrow _e_ in Fig. 115. It may be
designated as the femoral lever.

The plane of direction in which the femur, as seen by the rotation
just mentioned, is moved, exactly coincides in insects with that of
the tibia and the foot, while all can be simultaneously raised or
dropped, or, as the case may be, stretched out or retracted.
Therein, therefore, lies an essential difference from the fully
developed extremities of vertebrates among which, even on the lever
arms which are stationary at the end, an extensive turning is
possible.

The muscles which move the tibia, and indirectly the femur, also
consist of an extensor muscle which is situated in the upper side of
the femur (Fig. 116, _s_, Fig. 115, _f_), and of a flexor (Fig. 116,
_b_, Fig. 115, _g_), which lies under the former.

The stilt-like spines on the point (Figs. 115 and 118, _L_{3}n_) on
which this segment is directly supported are important parts of the
tibia. (Graber.)

[Illustration:

FIG. 117.—Left fore leg of a cerambycid beetle: _h_, coxa; _r_,
trochanter; _o_, femur; _u_, tibia; _f_, tarsus; _k_, claw.—After
Graber.
]

Considering the respective positions of the individual levers of the leg
and the nature of the materials of which they are made, the legs of
insects may be likened, as Graber states, to elastic bows, which, when
pressed down together from above, their own indwelling elasticity is
able to raise again and thus keep the body upright.

This is very plainly shown in certain stilt-legged bark-beetles, in
which, as in a rubber doll, as soon as the body is pressed down on the
ground, the organs of motion extend again without the intervention of
muscles; indeed this experiment succeeds even with dead, but not yet
wholly stiff, insects.

Graber then turns to the analysis of the movements of insect legs when
in motion, and the mode of walking of these insects in general. This
subject had been but slightly investigated until Graber made a series of
observations and experiments, of which we can give only the most
important results.

The locomotion of insects is an extremely complicated subject.

Let us consider, Graber says, first, a running or carabid beetle,
when walking merely with the fore and hind legs. The former will be
bent forward and the latter backward.

“Let us begin with the left fore leg (Fig. 118, _L_{1}_). Let the
same be extended and fixed on the ground by means of its sharp claws
and its pointed heel. Now what happens when the tibial flexors draw
together? As the foot, and therefore the tibia also, have a firm
position, then the contraction of the muscles named must cause the
femur to approach the tibia, whereby the whole body is drawn along
with it. This individual act of motion may be well studied in
grasshoppers when they are climbing on a twig by stretching out
their long fore leg directly forward, and then drawing up the body
through the shortening of the tibial flexors until the middle leg
also reaches the branch.

“But while the fore legs advance the body by drawing the free lever
to the fixed leg-segment, the hind legs do this in exactly the
opposite way. The hind leg, namely, seeks to stretch out the tibia,
and thus to increase the angle of the knee (_R_{3})_, thereby giving
a push on the ground, by means of which the body is shoved forward a
bit.

“Though it might be supposed that the feet would remain stationary
during the extension or retraction of the limbs, this never occurs
in actual walking. Not merely the upper, but also the lower, thigh
is either drawn in or stretched out, as the case may be. The latter
then describes a straight line with its point during this scraping
or scratching motion (Fig. 115, _no_), which is obviously the chord
to that quadrant which would be drawn by the tibia or foot in a
yielding medium, as water, for instance. But even this motion
results extremely rarely, and never in actual walking. If we fix our
eye anew upon the fore leg at the very moment when it is again
retracted, after the resultant ‘fixing,’ we shall then observe that
the hip also is simultaneously turned backward in a definite angle.
The tibia would describe the arc _nq_ (Fig. 115) by means of the
latter alone.

“This plane, in conjunction with the rectilinear ‘movement’ (_no_)
obtained by the retraction of the tibia, produces a path (_nr_), and
this is what is actually described by a painted foot upon a properly
prepared surface, as a sheet of paper;[22] supposing, however, that
the body in the meantime is not moved forward by other forces. In
the last case, and this indeed always takes place in running, the
trunk is moved a bit forward, together with the leg which is just
describing its curve with a rapidity corresponding to the momentum
obtained; the result of this is that the curve of the foot from its
beginning (_n_) to its end (_a_) bends round close to itself, just
as a man who, when on board a ship in motion, walks across it
diagonally, and yet on the whole moves forward, because his line of
march, uniting with that of the ship, results in a change of
position in space.

“The case is the same in the middle and hind legs, which must make a
double course also, yet in such a way that the straight line is
drawn, not during the retraction, but during the extension; during
which, however, quite as in the fore leg, the members mentioned
(_R_{3}_) gradually approach the body.

“When the legs have reached the maximum of their retraction, or of
their extension, as the case may be, and therefore the end of their
active course for that time, then begins the opposite or backward
movement; that is, the fore legs are again extended, while their
levers draw the remaining legs together again.

[Illustration:

FIG. 118.—A Carabus beetle in the act of walking or running: three
legs (_L_{1}_, _R_{2}_, _L_{3}_) are directed forward, while the
others (_R_{1}_, _L_{2}_, _R_{3}_), which are directed backward
toward the tail, have ended their activity; _ab_, _cd_, and _ef_
are curves described by the end of the tibiæ, and passing back to
the end of the body; _bh_, _di_, and _fg_ are curves described by
the same legs during their passive change of position.—After
Graber.
]

“At the same time, as we may see by the uniting leg, the limb is
either a little raised, that there may be no unnecessary friction,
or it remains during the passive step also, with its means of
locomotion in slight contact with the ground.

“The curve of two steps, as inscribed by the end of the tibia of the
left fore leg of a stag-beetle, affords an instructive summary of
the conditions of which we have been speaking (Fig. 121, _B_). We
see two curves. The thick one (_ab_), directed toward the axis of
the body, corresponds to the effective act of a single walking
function, which brings the body a bit forward; the thinner, on the
other hand, or we might say the hair line (_bc_), which, however, is
but rarely made quite clearly, is produced by the ineffectual
backward movement, by which the insect again approaches its working
posture (_c_). It is at first placed at some distance from the body,
in order that (like _c_ also) it may draw near to the body again;
but in such a way, naturally, that it coincides with the
starting-point of the following active curve (_cd_). It is evident
that even the passive curve is not the imprint of the movement
accomplished exclusively by the leg, for this latter, while
struggling to reach its resting-place, is really involuntarily
carried forward with the rest of the body.

“The scroll-like lines drawn by the swimming beetle (Dyticus), with
the large, sharp points of its hind tibia, are also very instructive
(Fig. 119, _A_).

[Illustration:

FIG. 119.—_A_, trail curves described by the tibial spines of the
right and left hind limb of Dyticus. _B_, the same made by the
right hind leg (_r_{3}_) alone. Natural size.—After Graber.
]

[Illustration:

FIG. 120.—The same by the two hind legs of Melolontha: _a_, the
active and thickened section of the curve. Natural size.
]

[Illustration:

FIG. 121.—_A_, track curves of two of the tibial spines of the left,
middle legs of a stag-beetle. Natural size. _B_, the same
enlarged; _fg_, the longitudinal axis of the trunk; _cd_ and _ab_,
the active curve passing inward,—_bc_ and _de_, the passive going
outward. _C_, two curves described by the left hind legs; in this
case, the curves are not inwards or backwards, but partly directly
inward (_b_), and in part obliquely forwards (_a_).
]

“The diversions and modifications in the course of the active step,
as furnished by the moving factor of the remaining legs, are already
clearly illustrated by the curves shown by the joints of the hind
tibia of a May-beetle (Fig. 120) and a stag-beetle (Fig. 121, _c_).
The actual faint line in this case does not run from the front
toward the back, as would correspond to the active leg-motion, but
either directly inward (Fig. 121, _cb_), or even somewhat to the
front. In the May-beetles, and even more in the running
garden-beetle, the curves of the hind legs present themselves as
screw-like lines (Fig. 122, _l_{3}_), while the scrawling of the
remaining members (_l_{1}_, _l_{2}_) is much simpler.

“Inasmuch as we now have a cursory knowledge of the movements made
by each individual leg for itself,—movements, however, which plainly
occur very differently according to the structure of these
appendages,—the question now is of the combined play, the total
effect of all the legs taken together, and therefore of the walk and
measure of the united work of the foot.

“In opposition to the caterpillars and many other crawling animals
which extend their legs in pairs and really swing them by the
worm-like mode of contraction of the dermo-muscular tube, the legs
of fully grown insects are moved in the contrary direction and in no
sense in pairs, but alternately—or, more strictly speaking, in a
diagonal direction.

“For an examination of the gait of insects, we choose, for obvious
reasons, those which have very long legs and which at the same time
are slow walkers.

“Insects may be called ‘double-three-footed,’ from the manner in
which they alternately place their legs. There are always three legs
set in motion at the same time, or nearly so, while in the meantime
the remaining legs support the body, after which they change places.

[Illustration:

FIG. 122.—The same by the left fore (_l_{1}_), middle (_l_{2}_), and
hind, leg (_l_{3}_) of a Carabus. Natural size.
]

[Illustration:

FIG. 123.-Tracks of a _Blaps mortisaga_ marked by the differently
painted tibial points: ●, tracks of fore, —○, middle, —/, hind
leg. Natural size.
]

[Illustration:

FIG. 124.—Tracks of _Necrophorus vespilio_. Natural size.
]

“To be more exact, it is usually thus: At first (Fig. 118) the left
fore leg (_L_{1}_) steps out, then follows the right middle leg
(_R_{2}_), and the left hind leg (_L_{3}_). Then while the left fore
leg begins to retract and thus make the backward movement, the right
fore leg is extended, whereupon the left middle leg and the right
hind leg are raised in the same order as the first three feet.”

Graber[23] painted the feet of beetles and let them run over paper,
and goes on to say:

“Let us first pursue the tracks of the Blaps, for example (Fig.
123). Let the insect begin its motion. The left fore leg stands at
_a_, the right middle leg at β, and the left hind leg at _c_. The
corresponding number of the other set of three feet at α, _b_, γ. At
the first step the three feet first mentioned advance to _a′_β′_c′_,
the second set on the other hand to α′_b′_γ′. Thereby the tracks
made by the successive steps fall quite, or almost quite, on each
other, as appear also in the tracks of a burying beetle (Fig. 124).

“As the fore legs are directed forward and the hind legs backward,
while the middle legs are placed obliquely, the reason of the more
marked impressions of the latter is evident.

“The highest testimony to the precise exactitude and accuracy of the
walking mechanism of insects is furnished by the fact that in most
insects, and particularly in those most fleet of foot, which,
whether they are running away or chasing their prey, must be able to
rely entirely upon their means of locomotion;—the fact, we say, that
whether they desire to move slowly or more quickly, the distances of
the steps, measured by the length as well as by the cross-direction,
hardly differ a hair’s breadth from one another, and this is also
the case when the tarsi are cut off and the insects are obliged to
run on the points of their heels (tibiæ).

“Thence, inasmuch as the trunk of insects is carried by two legs and
by one on each side alternately, it may surely be concluded _a
priori_ that when walking it is inclined now to the right and now to
the left, and that the track, too, which is left behind by a precise
point of the leg, can in no wise be a straight line; and in reality
this is not the case.

“A plainly marked regular curve, which approaches a sinuous line, as
seen in Fig. 125, is often obtained by painting many insects, for
example Trichodes, Meloë, etc., which, when running, either bring
the end of their hind body near to the ground or into contact with
it.

[Illustration:

FIG. 125.—Tracks of Trichodes; the middle sinuous line is made by
the tip of the abdomen. Natural size.
]

[Illustration:

FIG. 126.—Tracks of another insect which, in running, can only use
three legs (_r_{1}_, _l_{4}_, _r_{3}_) which become indicated
differently from normal conditions. Natural size.
]

[Illustration:

FIG. 127.—The same of an insect crossing over a surface inclined 30°
from the horizon, whereby the placing of the feet becomes changed.
Natural size.—This and Figs. 120–126 after Graber.
]

“The locomotive machine of insects may be called, to a certain
extent, a double set of three feet each, as most insects, and
particularly those provided with a broad trunk, are able to balance
themselves with one of these two sets of feet, and indeed when
walking, as well as when standing still, can move about even better
with one set of these feet than with four legs. In the latter case,
that is, if one cuts off a pair of legs from an insect, the trunk
can balance itself only with extreme difficulty, and there is
therefore little prospect that insects will ever become four-footed.

“But if one compels insects to run on three legs, he will thus make
the interesting discovery that to make up the deficiency they place
the remaining feet and bring them to the ground somewhat differently
than when the second set of feet is active. Figs. 124 and 126 may be
compared for this purpose. The former shows the footprints of a
burying beetle running with all six legs, the latter the track of
the same insect, which, however, has at its disposal only the right
fore leg, the left middle leg, and the right hind leg. One may
plainly see here that the track of the hind leg on the right side
(_r_{3}_) approaches the track of the middle leg on the left side,
and then further, that the _right fore leg_ (_r_{1}_) _steps out
more to the right to make up for the deficiency of the middle leg_.

“A similar adaptation of the position of the legs, which is entirely
dependent on the choice of the insect, may also be observed there,
if one compels insects which are not provided with corresponding
adhesive lobes to run away over crooked surfaces. Fig. 123 shows the
footprints of a Blaps when running upon a horizontal plane. Fig.
127, on the contrary, shows the tracks of the legs when going
diagonally over a gradually inclined surface. Here, also, the insect
holds on with his fore and middle legs (_r_{1}_, _r_{2}_) stretched
upward, whereby also the impressions on both sides come to lie
farther apart than in the normal mode of walking.

“It will not surprise the reader who is familiar with the gait of
crabs, to hear that many insects also understand the laudable art of
going backward, wherein the hind legs simply change places with the
fore legs.

“The jumping motion of insects may be best studied in grasshoppers.
When these insects are preparing for a jump, they stretch out the
upper thigh horizontally, clap the tibiæ together, and also retract
the foot-segment. After a slight pause for rest, during which they
are getting ready for the jump, they then jerk the tibiæ suddenly
backward and against the ground with all their strength by means of
the extensor muscles.”

The correctness of Graber’s views has been confirmed by Marey by
instantaneous photographs (Figs. 128, 129).

=Locomotion on smooth surfaces.=—How flies and other insects are able to
walk up, or run with the body inverted, on hard surfaces has been lately
discovered by Dewitz, Dahl, and others. All authors are agreed that this
power is due to the presence of the specialized empodium of each tarsus.

Dewitz confirmed the opinion of Blackwell, that a glutinous liquid is
exuded from the apices of the tenent hairs which fringe the empodium. By
fastening insects feet uppermost on the under side of a covering glass
which projects from a glass slide, the hairs which clothe the empodia of
the foot of a fly (_Musca erythrocephala_) may be seen to be tipped with
drops of transparent liquid. On the leg being drawn back from the glass,
a transparent thread is drawn out, and drops are found to be left on the
glass. In cases where these hairs are wanting, as in the Hemiptera, the
adhesive fluid exudes directly from pores in the foot. In the beetles
(_Telephorus dispar_) and other insects the tenent hairs on the foot end
in sharp points, below which are placed the openings of the canals. The
glands, Dewitz states, are chiefly flask-shaped and unicellular,
situated in the hypodermis of the chitinous coat; each gland opening
into one of the hairs (Fig. 108); they are each invested by a
structureless tunica propria, and contain granular protoplasm, a nucleus
placed at the inner side, and a vesicle, prolonged into a tube which,
traversing the neck of the gland, is attached to the root of the hair;
the vesicle receiving the secretion. Each gland is connected with a fine
nerve-twig, and secretion is probably voluntary. Among the tenent hairs
of the empodium are others which must be supplied with a nerve, forming
tactile hairs, as they each proceed from a unicellular ganglion (Fig.
108, _n″_). The secretion is forced out of the gland by the contraction
of the protoplasm, Dewitz having seen the secretion driven out from the
internal vesicle into its neck.

[Illustration:

FIG. 128.—The walk of an orthopterous insect: series to be followed
from right to left.—After Marey.
]

[Illustration:

FIG. 129.—Beetle walking: series to be followed from left to
right.—After Marey.
]

[Illustration:

FIG. 130.—_A_, end of an adhesive hair of a weevil (Eupolus): _i′_,
canal: _i‴_, its external opening at the end of the hair. _B_, end
of a similar hair of Telephorus with drops of the secretion.—After
Dewitz.
]

In the spherical last tarsal joint of Orthoptera (Fig. 109), which
is without these tenent hairs, nearly all the cells of the
hypodermis are converted into unicellular glands, each of which
sends out a long, fine, chitinous tubule, which is connected with
its fellows by very fine hairs and is continuous with the chitinous
coat of the foot and opens through it. The sole of the foot is
elastic and adapts itself to minute inequalities of surfaces, while
the anterior of each tarsal joint is almost entirely occupied by an
enlargement of the trachea, which acts on the elastic sole like an
air chamber, rendering it tense and at the same time pliant. Dewitz
adds that the apparatus situated on the front legs of the male of
_Stenobothrus sibiricus_ (Fig. 131) must have the function of
causing the legs to adhere closely to the female by the excretion of
an adhesive material. The hairs of the anterior tarsi of male Carabi
also appear to possess the power of adhesion. In the house-fly the
empodia seem to be only called into action when the insect has to
walk on vertical smooth surfaces, as at other times they hang
loosely down.

Burmeister observed the use of a glutinous secretion for walking in
dipterous larvæ, and Dewitz found that the larva of a Musca used for
this purpose a liquid ejected from the mouth. The larvæ of another
fly (_Leucopis puncticornis_) perform their loop-like walk by
emitting a fluid from both mouth and anus. A Cecidomyia larva is
able to leap by fixing its anterior end by means of an adhesive
fluid. The larva of the leaf-beetle, Galeruca, moves by drawing up
its hinder end, fixing it thus, and carrying the anterior part of
the body forward with its feet until fully extended, when it breaks
the glutinous adhesion. The abdominal legs of some saw-fly larvæ
have the same power.

Dahl could not detect in the foot of the hornet (_Vespa crabro_) any
space which could be considered as a vacuum.

[Illustration:

FIG. 131.—_Stenobothrus sibiricus_ pairing: _A_. the ♂, fore tarsus
(_t_) greatly enlarged; _ar_, arolia; _p_, pulvillus.—After
Pagenstecher.
]

Simmermacher states that in most cases of climbing beetles the tubular
tenent hairs pour out a secretion (Figs. 133, 134), “and it is probable
that we have here to do with the phenomena not of actual attachment by,
as it were, gluing, but of adhesion; the orifice of the tubes is divided
obliquely, and the tubes are, at this point, extremely delicate and
flexible, so as to adhere by their lower surface; in this adhesion they
are aided by the secreted fluid.” In the case of the Diptera he does not
accept the theory by which the movement of the fly along smooth surfaces
is ascribed to an alternate fixation and separation, but believes in a
process of adhesion, aided by a secretion, as in many Coleoptera. (In
the Cerambycidæ there is no secretion, and the tubules are merely
sucking organs, like those observed in the male Silphidæ.) “The
attaching lobes, closely beset with chitinous hairs, are enabled, in
consequence of the pressure of the foot, to completely lie along any
smooth surface; this expels the air beneath the lobes, which are then
acted on by the pressure of the outer air.” (Journ. Roy. Micr. Soc.,
1884, p. 736.) Another writer (Rombouts) thinks this power is due to
capillary adhesion.

[Illustration:

FIG. 132.—Fore leg of ♂ Dyticus, under side, with sucker, formed of 3
enlarged tarsal joints: with a small cupule highly magnified. ×
120.—After Miall.
]

The action of the pulvillus and claws when at rest or in use by the
honey-bee is well shown by Cheshire (Fig. 135, _B_). In ascending a
rough surface, “the points of the claws catch (as at _B_) and the
pulvillus is saved from any contact, but if the surface be smooth, so
that the claws get no grip, they slide back and are drawn beneath the
foot (as at _A_), which change of position applies the pulvillus, so
that it immediately clings. It is the character of the surface, then,
and not the will of the bee, that determines whether claw or pulvillus
shall be used in sustaining it. But another contrivance, equally
beautiful, remains to be noticed. The pulvillus is carried folded in the
middle (as at _C_, Fig. 105), but opens out when applied to a surface;
for it has at its upper part an elastic and curved rod (_cr_, Figs. 105
and 135), which straightens as the pulvillus is pressed down; _C_ and
_D_, Fig. 135, making this clear. The flattened-out pulvillus thus holds
strongly while pulled, by the weight of the bee, along the surface, to
which it adheres, but comes up at once if lifted and rolled off from its
opposite sides, just as we should pull a wet postage stamp from an
envelope. The bee, then, is held securely till it attempts to lift the
leg, when it is freed at once; and, by this exquisite yet simple plan,
it can fix and release each foot at least twenty times per second.”
(Bees and Bee-keeping, p. 127.)

[Illustration:

FIG. 133.—Cross-section through a tarsal joint of fore leg of Dyticus,
♂, showing the stalked chitinous suckers (_s_), with a marginal
bristle on each side: _t_, trachea; _a_, an isolated tubule or
sucker of Loricera,—_b_, of Chlænius,—_c_, of Cicindela; _d_, two
views of one of _Necrophorus germanicus_, ♂.
]

[Illustration:

FIG. 134.—Section through the tarsus of a Staphylinid beetle; the
glandular or tenent hairs arising from chitinous processes. _A_,
section through the tarsal joint of the pine weevil, _Hylobius
abietis_, showing the crowded, bulbous, glandular, or tenent hairs
arising from unicellular glands.—This and Fig. 133 after
Simmermacher.
]

Ockler divides the normal two-clawed foot into three subtypes: (1) with
an unpaired median empodium; (2) with two outer lateral adhesive lobes;
(3) with two adhesive lobes below the claws; the latter is the chief
type and forms either a climbing or a clasping foot. The amount of
movement possessed by the claws is limited, and what there is, is
effected by means of an elastic membrane and the extensor plate (Fig.
110). The “extensor sole” which is always present in insects with an
unpaired median fixing or adhesive organ (empodium) is to be regarded as
a modification of the extensor seta. The extensor plate is peculiar to
an insect’s foot. Ockler states that the so-called “pressure plate” of
Dahl is only a movably articulated, skeletal, supporting plate for the
median fixing lobule.

[Illustration:

FIG. 135.—Honey-bee’s foot in the act of climbing, showing the
automatic action of the pulvillus, × 30: _A_, position of foot in
climbing on a slippery surface, or glass; _pv_, pulvillus; _fh_,
tactile hairs; _un_, unguis; _t_, last tarsal joint. _B_, position
of foot in climbing rough surface. _C_, section of pulvillus just
touching flat surface; _cr_, curved rod. _D_, the same applied to
the surface.—After Cheshire.
]

=Climbing.=—In certain respects the power of climbing supplies the want
of wings, and even exists often in house-flies among which there is
shown a many-sided motion that is quite unheard of in other groups of
insects.

The best climbers are obviously those insects which live on trees and
bushes, as, for example, longicorn beetles and grasshoppers. These may
be accurately called the monkeys of the insect kind, even if their
movements take place less gracefully, and indeed rather stiffly and
woodenly. We already know what are the proper climbing organs; that is,
the sharp easily movable claws on the foot. With the help of these claws
certain insects, May-beetles for example, can hang upon one another like
a chain; indeed, bees and ants in this manner bind themselves together
into living garlands and bridges. There are still added to the chitinous
hooks flaps and balls of a sticky nature, by help of which likewise the
insects glue themselves together. To facilitate the spanning of still
thicker twigs, the climbing foot of insects has a greater movability
even than when it only serves as a sole. (Graber.)

=The mode of swimming of insects.=—To study the swimming movements of
insects, let us examine a Dyticus. It will appear, as Graber states, to
be wonderfully adapted to its element.

“The body resembles a boat. There is nowhere a projecting point or a
sharp corner which would offer unnecessary resistance to motion;
bulging out in the middle and pointed at the end, it cuts through
the resistance of the water like a wedge. The movable parts, the
oars, seem to be as well fitted for their purpose as the burden to
be moved by them. That the hind legs must bear the brunt of this
follows from their position exactly in the middle of the body, where
it is widest. In other insects also these legs are used for the same
purpose as soon as the insects are put in the water. But the
swimming legs of water-beetles are oars of quite peculiar
construction. _They are not turned about in the coxæ, as are other
legs, but at the foot-joint._ The coxa, namely, has grown entirely
together with the thoracic partition. The muscles we have mentioned,
exceeding in strength all the soft parts taken together, take hold
directly of the large wing-shaped tendons of the upper thigh, and
extend and retract the leg in one of the planes lying close to the
abdominal partition. The foot forms the oar, however. It is very
much lengthened and still more widened, and can be turned and bent
in by separate muscles in such a way that in the passive movement,
that is, the retraction, the narrow edge is turned to the fore, and
therefore to the medium to be dislodged; however, as soon as the
active push is to be performed and the leg is extended with greater
force, it cuts down through the water with its whole width. These
effective oar-blades are still considerably enlarged by the hairs
arising on the side of the foot, which spread out at the decisive
moment.

“Every one knows that the oar-blades of swimming beetles always go
up and down simultaneously and in regular time. On the other hand,
as soon as one puts a Dyticus on the dry land, _i.e._ on an
unyielding medium, it uses its hind legs entirely after the manner
of other land insects; that is, they are drawn in and extended again
_alternately_, as takes place clearly enough from the footsteps in
Fig. 119, _A_. We learn from this that water insects have not yet,
from want of practice, forgotten the mode of walking of land
insects.

“The forcing up of the water as a propelling power is added to the
repulsion produced by the strong strokes of the oars. If the beetle
stood up horizontally in the water, he would be lifted up.

“As the trunk, however, assumes an oblique position when the insect
wishes to swim, one can then imagine the driving up of the water as
being divided into two forces, one of which drives the body forward
in a horizontal direction, while the other, that is, the vertical
component, is supplied by the moving of the oars. The swimming
insect is thus, as it were, a snake flying in the water.

“The long streamer-like hind legs of many water-bugs, for example
Notonecta, approach more nearly our artificial oars. These legs are
turned out from the bottom.

“There is no doubt but that the legs of insects, as regards the
many-sidedness and exactitude of their locomotive actions, place the
similar contrivances of other animals far in the shade. We shall be
forced to admire these ingenious levers still more, however, when we
take into consideration their energy and strength. That the force
with which the locomotive muscles of insects is drawn together is
enormous compared with that of vertebrates, we may learn if we try
to subdue the rhythmical movements of the thorax of a large
butterfly by the pressure of our finger or to open against the
insect’s will the closed jumping leg of a grasshopper, or the
fossorial shovel of a mole-cricket.”

LITERATURE ON LEGS AND FEET

=MacLeay, W. S.= On the structure of the tarsus in the tetramerous
Coleoptera of the French entomologists. (Trans. Linn. Soc. London,
xv, 1825, pp. 63–73.)

=Speyer, O.= Untersuchung der Beine der Schmetterlinge. (Isis, 1843,
pp. 161–207, 243–264.)

=Pokorsky Joravko, A. von.= Quelques remarques sur le dernier article
du tarse des Hyménoptères. (Bull. Soc. imp. Natur. Moscou, 1844,
xvii, pp. 140–159. Ref. in Isis, 1848, v, p. 347.)

=Rossmassler, E. A.= Das Bein der Insekten. (Aus der Heimath, 1860, 3
kap., pp. 327–334, Fig.)

=West, Tuffen.= The foot of the fly; its structure and action;
elucidated by comparison with the feet of other insects, etc. Part
I. (Trans. Linn. Soc. London, xxiii, 1861, pp. 393–421, 1 Pl.)

=Sundevall, C.= On insektenas extremiteter samt deras hufoud och
munddelar. (Kongl. Vetenskaps Akad. Handlingar. iii, Nr. 9, 1861.)

=Lindemann, C.= Notizen zur Lehre vom ausseren Skelete der Insekten
(Gelenke und Muskeln der Füsse). 1 Taf. (Bull. Soc. imp. d. Natur.
Moscou, xxxvii, 1864, pp. 426–432.)

=Liebe, O.= Die Gelenke der Insekten. Chemnitz, 1873. 4º. 1 Taf.

=Canestrini, J.= Ueber ein sonderbares Organ der Hymenopteren. (Zool.
Anzeiger, 1880, pp. 421, 422.)

=Dahl, F.= Beiträge zur Kenntnis des Baues und der Funktionen der
Insektenbeine. (Archiv f. Naturgesch. 1 Jahrg., 1884, pp. 146–193, 3
Taf. Sep., 48 pp. Vorlauf. Mitteil, in Zool. Anz., 1884, pp. 38–41.)

=Langer, K.= Ueber den Gelenkbau bei den Arthrozoen. Vierter Beitrag
zur vergleichenden Anatomie und Mechanik der Gelenke. (Denkschriften
der Akad. d. Wissensch. Wien, xviii, Bd. Physikal.-mathem. Classe,
pp. 99–140. 3 Taf.)

=Graber, Vitus.= Ueber die Mechanik des Insektenkörpers. (Biolog.
Centralbl., iv, 1884, pp. 560–570.)

—— Die ausseren mechanischen Werkzeuge der Tiere, ii Teil. Wirbellose
Tiere, 1886, pp. 175–182, 208–210.

=Dewitz, H.= Ueber die Fortbewegung der Tiere an senkrechten glatten
Flächen vermittelst eines Sekretes. 3 Taf. (Pflüger’s Archiv f. d.
ges. Physiologie, xxxiii, 1884, pp. 440–481.)

=Ockler, A.= Das Krallenglied am Insektenfuss. (Archiv f. Naturgesch.,
1890, pp. 221–262, 2 Taf.)

LITERATURE OF LOCOMOTION (WALKING, ETC.)

=Carlet, G.= Sur le mode de locomotion des chenilles. (Compt, rend.
Acad. Paris, 1888, cvii, pp. 131–134. Naturwiss. Rundschau, iii
Jahrg., 1888, No. 42, p. 543.)

—— De la marche d’un insecte rendu tetrapode par la suppression d’une
paire de pattes. (Ibid., pp. 565, 566.)

—— Sur la locomotion des insectes et des arachnides. (Ibid., 1879, T.
89, pp. 1124, 1125.)

—— Ueber den Gang eines vierfüssig gemachten Insekts. (Naturwiss.
Rundschau, viii Jahrg., 1888, pp. 666–667; Compt. rend. 1888, cvii.)

=Demoor, J.= Recherches sur la marche des insectes et des arachnides.
Étude experimentale d’Anatomie et de Physiologie comparées. (Archiv
de Biologie, Liège, 1880, 42 pp. 3 Pls.)

—— Ueber das Gehen der Arthropoden mit Berücksichtigung der
Schwankungen des Körpers. (Compt. rend. Acad. d. Sc. Paris, 1890,
cxi, pp. 839–840.)

=Osten-Sacken, C. R. von.= Ueber das Betragen des kalifornischen
flügellosen Bittacus (apterus McLachl.). (Wiener Ent. Zeit., 1882,
pp. 123.)

=Dixon, H. H.= Preliminary note on the walking of some of the
Arthropoda. (Proc. R. Dublin Soc. vii, pp. 574–578, 1892. Also
Nature, 1897.)

Also the works of Graber, Marey, Cheshire, etc.

LITERATURE OF WALKING ON SMOOTH SURFACES

=Blackwell, J.= Remarks on the pulvilli of insects. (Trans. Linn. Soc.
London, xvi, 1831, pp. 487–492, 767–770.)

=Lowne, B. T.= On the so-called suckers of Dytiscus and the pulvilli
of insects. (Trans. Roy. Micr. Soc., pp. 267–271, 1871, 1 Pl.)

=West, Tuffen.= On certain appendages to the feet of insects
subservient to holding or climbing. (Journ. of the Proceed. Linn.
Soc. London, Zoölogy, vi, 1862, pp. 26–88.)

=Dewitz, H.= Ueber die Fortbewegung der Tiere an senkrechten, glatten
Flächen vermittelst eines Sekrets. (Pflüger’s Archiv f. d. ges.
Physiologie, xxxiii, 1884, pp. 440–481. 3 Taf. Also Zool. Anzeiger,
1884, pp. 400–405.)

—— Wie ist es den Stubenfliegen und anderen Insekten möglich, an
senkrechten Glaswanden emporzulaufen. (Sitzungsb. Ges. naturf.
Freunde zu Berlin, 1882, pp. 5–7.)

—— Weitere Mitteilungen über den Klettern der Insekten (Ibid., 1882,
pp. 109–113).

—— Die Befestigung durch einen klebenden schleim beim springen gegen
senkrechte Flächen. (Zool. Anzeiger, 1883, pp. 273, 274.)

—— Ueber die Wirkung der Haftlappchen toter Fliegen. (Ent. Nachr., x
Jahrg., 1884, pp. 286, 287.)

—— Weitere Mitteilungen über das Klettern der Insekten an glatten
senkrechten Flächen. (Zoolog. Anzeiger, 1885. viii Jahrg., pp.
157–159.)

—— Richtigstellung der behauptungen des Herrn F. Dahl. (Archiv f.
mikroskop. Anat., 1885, xxvi, pp. 125–128.)

=Rombouts, J. E.= Ueber die Fortbewegung der Fliegen an glatten
Flächen. (Zool. Anzeiger, 1884, pp. 619–623.)

—— De la faculté qu’out les mouches de se mouvoir sur le verre et sur
les autres corps polis. (Archiv Museum Teyler (2), 4 Part, pp. 16.
Fig.)

=Simmermacher, G.= Untersuchungen über Haftapparate an Tarsalgliedern
von Insekten. (Zeitschr. f. wissensch. Zool. xl, 1884, pp. 481–556.
3 Taf., 2 Figs. Also Zoolog. Anzeiger, vii Jahrg., 1884, pp.
225–228.)

—— Antwort an Herrn Dr. H. Dewitz. (Ibid., pp. 513–517.)

=Dahl, F.= Die Fussdrüsen der Insekten. (Archiv f. mikroskop. Anat.,
1885, xxv, pp. 236–263. 2 Taf. See also p. 118.)

=Emery, C.= Fortbewegung von Tieren an senkrechten und überhangenden
glatten Flächen. (Biolog. Centralbl., 1884, 4 Bd., pp. 438–443.)

=Léon, N.= Disposition anatomique des organes de succion chez les
Hydrocores et les Géocores. (Bull. Soc. des Medec. et Natur, de
Jassy., 1888.)

_d._ The wings and their structure

The insects differ from all other animals except birds in possessing
wings, and as we at the outset have claimed, it is evidently owing to
them that insects are numerically so superior to any other class of
animals, since their power of flight enables them to live in the air out
of reach of many of their enemies, the greatest destruction to insect
life occurring in the wingless larval and pupal stages.

The presence of wings has exerted a profound influence on the shape and
structure of the body, and it is apparently due to their existence that
the body is so distinctly triregional, since this feature is least
marked in the synapterous insects. The wings are thin, broad leaf-like
folds of the integument, attached to the thorax and moved by powerful
muscles which occupy the greater part of the thoracic cavity. The two
pairs of wings are outgrowths of the middle and hinder part of the
thorax, the anterior pair being attached to the mesothoracic and the
hinder pair to the metathoracic segment. The larger pair is developed
from the middle segment of the thorax. The differentiation of the
tergites into scutum, scutellum, etc., is the result of the appearance
of wings, because these sclerites are more or less reduced or effaced in
wingless insects, such as apterous Orthoptera and moths, ants, etc.

The size of the hinder thoracic segments is closely related to that of
the wings they bear. In those Orthoptera which have hind wings larger
than those of the fore pair, the metathorax is larger than the
mesothorax. In such Neuroptera as have the hind wings nearly or quite as
large as the anterior pair, or in the Trichoptera and in the Hepialidæ,
the metathorax is nearly as large as the mesothorax, while in Coleoptera
the metathorax is as large and often much larger. In the Ephemeridæ,
Diptera, and Hymenoptera, which have either only rudimentary (halteres)
or small hind wings, the metathorax is correspondingly reduced in size.

The wings morphologically, as their development shows, are simple
sac-like outgrowths of the integument, _i.e._ of the free hinder edge of
the tergal plates, their place of origin being apparently above the
upper edge of the epimera or pleural sclerites. Calvert[24] however,
regards the upper lamina of the wing as tergal, and the lower, pleural.

The wings in most insects are attached to the thorax by a membrane
containing several little plates of chitin called by Audouin
articulatory epidemes.

The wings, then, are simple, very thin chitinous lamellate expansions of
the integument, which are supported and strengthened by an internal
framework of hollow chitinous tubes.

=The veins.=—The so-called “veins” or “nervures,” which are situated
between the upper and under layers of the wing are so disposed as to
give the greatest lightness and strength to the wings. Hagen has shown
that in the freshly formed wings these two layers can be separated, when
it can be seen that the veins pass through each layer.

These veins are in reality quite complex, consisting of a minute central
trachea enclosed within a larger tube which at the instant the insect
emerges from the nymph, or pupa, as the case may be, is filled with
blood (Fig. 136). Since these tubes at first contain blood, which has
been observed to circulate through them, and since the heart can be most
easily injected through them, they may more properly be called veins
than nervures. The shape and venation of the wings afford excellent
ordinal as well as family and generic characters, while they also enable
the systematist to exactly locate the spots and other markings of the
wings. The spaces enclosed by the veins and their cross-branches are
called cells, and their shape often affords valuable generic and
specific characters.

[Illustration:

FIG. 136.—Cross-section of wing of Pronuba.—After Spuler.
]

[Illustration:

FIG. 137.—Cross-section of wing of Pieris: _s_, insertions of
scales.-After Spuler.
]

The structure of a complete vein is described by Spuler. In a
cross-section of a noctuid moth (_Triphæna pronuba_, Fig. 136) the
chitinous walls are seen to consist of two layers, an outer (U) and
inner (_c_), the latter of which takes a stain and lies next to the
hypodermis (_hy_). In the cavity of the vein is the trachea (_tr_),
which shows more or less distinctly the so-called spiral thread; within
the cavity are also Semper’s “rib” (_r_) and blood-corpuscles (_bc_),
which proves that the blood circulates in the veins of the completely
formed wing, though this does not apply to all Lepidoptera with hard
mature wings. We have been able to observe the same structure in
sections of the wing of Zygæna.

A cross-section of a vein of _Pieris brassicæ_ shows that the large
trachea is first formed, and that it extends along the track between the
protoplasmic threads connecting the two hypodermal layers.

The main tracheæ throw off on both sides a number of secondary branches
showing at their end a cell with an intracellular tracheal structure;
these accessory tracheæ afterwards branch out. The accessory or
transverse tracheæ often disappear, though in some moths they remain
permanently. Fig. 137 _tr_{2}_ represents these secondary veins in the
edge of the fore wing of _Laverna vanella_, arising from a main trachea
(_tr_) passing through vein I (_v_), two of the twigs extending to the
centre, showing that the latter has no homology with a true vein. Only
rarely and in strongly developed thick folds are the transverse tracheæ
provided with a chitinous thickening, as for example in _Cossus
ligniperda_. Since from such accessory tracheæ the transverse veins in
lepidopterous wings are developed, we can recognize in them the
homologies of the net-veins in reticulated venations. There is no
sharply defined difference between reticulated and non-reticulated
venations; no genetic difference exists between the two kinds of
venation, since there occur true Blattidæ both with and without a
reticulated venation (Spuler).

In the fore wings of Odonata, Psocina, Mantispidæ, and most Hymenoptera
is an usually opaque colored area between the costal edge and the median
vein, called the _pterostigma_.

In shape the wings are either triangular or linear oval, and at the
front edge the main veins are closer together than elsewhere, thus
strengthening the wings and affording the greatest resistance to the air
in making the downward stroke during flight. It is noticeable that when
the veins are in part aborted from partial disuse of the wings, they
disappear first from the hinder and middle edge, those on the costal
region persisting. This is seen in the wings of Embiidæ (Oligotoma),
Cynipidæ, Proctotrupidæ, Chalcids, ants, etc.

The front edge of the wing is called the costal, its termination in the
outer angle of the wing is called the apex; the outer edge (termen) is
situated between the apex and the inner or anal angle, between which and
the base of the wing is the inner or internal edge.

While in Orthoptera, dragon-flies, Termitidæ, and Neuroptera the wings
are not attached to each other, in many Lepidoptera they are loosely
connected by the loop and frenulum, or in Hymenoptera by a series of
strong hooks. These hooks are arranged, says Newport, “in a slightly
twisted or spiral direction along the margin of the wing, so as to
resemble a screw, and when the wings are expanded attach themselves to a
little fold on the posterior margin of the anterior wing, along which
they play very freely when the wings are in motion, slipping to and fro
like the rings on the rod of a window curtain.”

At the base of the hind wings of Trichoptera and in the lepidopterous
Micropteryx there is an angular fold (_jugum_) at the base of each wing
(Fig. 138); that of the anterior wings is retained in Eriocephala and
Hepialidæ.

[Illustration:

FIG. 138.—Venation of fore and hind wings of _Micropteryx purpurella_:
_j_, jugum, on each wing; _d_, discal vein; the Roman numerals
indicate veins I.-VIII. and their branches.
]

In the wings of Orthoptera as well as other insects, the fore wings,
especially, are divided into three well-marked areas, the costal,
median, and internal; of these the median area is the largest, and in
grasshoppers and crickets is more or less modified to form the musical
apparatus, consisting of the drum-like resonant area, with the file or
bow.

=The squamæ.=—In the calyptrate Muscidæ, a large scale-like membranous
broad orbicular whitish process is situated beneath the base of the
wing, above the halter; (Fig. 94, 10 _sq._) it is either small or
wanting in the acalyptrate muscids. Kirby and Spence state that when the
insect is at rest the two divisions of this double lobe are folded over
each other, but are extended during flight. Their exact use is unknown.
Kolbe, following other German authors, considers the term _squama_ as
applicable to the whole structure, restricting the term _alula_ to the
other lobe-like division.

More recently (1890 and 1897) Osten-Sacken recommends “_squamæ_; in
the plural, as a designation for both of these organs taken
together; _squama_, in the singular, would mean the posterior squama
alone, and _antisquama_ the anterior squama alone;” the strip of
membrane running in some cases between them, or connecting the
squama with the scutellum, should be called the _post-alar
membrane_. By a mistake Loew, and others following him, used the
word _tegula_ for _squama_, but this term should be restricted to
the sclerite of the mesothorax previously so designated (Fig. 90,
_A_, _t_). The squama or its two subdivisions has also by various
authors been termed alula, calypta, squamula, lobulus, axillary
lobe, aileron, cuilleron, schuppen, and scale. (Berlin Ent.
Zeitschrift, xli, 1896, pp. 285–288, 328, 338.)

=The halteres.=—In the Diptera the hind wings are modified to form the
_halteres_ or balancers, which are present in all the species, even in
Nycteribia, but are absent in Braula.

Meinert finds structures in the Lepidoptera which he considers as
the homologues of the halteres of Diptera. “In the Noctuidæ,” he
remarks, “I find arising from the fourth thoracic segment (segment
médiaire), but covered by hair, an organ like the halter of
Diptera.” (Ent. Tidskrift., i, 1880, p. 168.) He gives no details.

In the Stylopidæ, on the contrary, the fore wings are reduced to little
narrow pads, while the hind wings are of great size.

The _thyridium_ is a whitish spot marking a break in the cubital vein of
the fore wing of Trichoptera; these minute thyridia occur in the fore
wings of the saw-flies; there is also an intercostal thyridium on the
costal part of the wings of Dermaptera.

The fore wings of Orthoptera are thicker than the hinder ones, and serve
to protect the hind-body when the wings are folded; they are sometimes
called _tegmina_. It is noteworthy, that, according to Scudder, in all
the paleozoic cockroaches the fore wings (tegmina) were as distinctly
veined as the hinder pair, “and could not in any sense be called
coriaceous.” (Pretertiary Insects of N. A., p. 39.) Scudder also
observes that in the paleozoic insects as a rule the fore and hind wings
were similar in shape and venation, “heterogeneity making its appearance
in mesozoic times.” In the heteropterous Hemiptera, also, the basal half
of the fore wings is thick and coriaceous or parchment-like, and also
protects the body when they are folded; these wings are called
_hemelytra_. In the Dermaptera the small short fore wings are thickened
and elytriform.

=The elytra.=—This thickening of the fore wings is carried out to its
fullest extent in the fore wings of beetles, where they form the
sheaths, shards, or _elytra_, under which the hind wings are folded. The
indexed costal edge is called the _epipleurum_, being wide in the
Tenebrionidæ. During flight “the elytra are opened so as to form an
angle with the body and admit of the free play of the wings” (Kirby and
Spence). In the running beetles (Carabidæ), also in the weevils and in
many Ptinidae, the hind wings are wanting, through disuse, and often the
elytra are firmly united, forming a single hard shell or case. The
firmness of the elytra is due both to the thickness of the chitinous
deposit and to the presence of minute chitinous rods or pillars
connecting the upper and lower chitinous surfaces.

[Illustration:

FIG. 139.—Longitudinal section through the edge of the elytrum of
_Lina ænea_: _gl_, glands; _r_, reservoir; _fb_, fat-body; _m_,
matrix; _u_, upper,—_l_, lower, lamella.—After Hoffbauer.
]

Hoffbauer finds that in the elytra of beetles of different families the
venation characteristic of the hind wings is wanting, the main tracheæ
being irregular or arranged in closely parallel longitudinal lines, and
nerve-fibres pass along near them, sense-organs being also present. The
fat-bodies in the cavity of the elytra, which is lined with a matrix
layer, besides nerves, tracheæ, and blood, contain secretory vesicles
filled with uric-acid concretions such as occur in the fat-body of
Lampyris. There are also a great many glands varying much in structure
and position, such occurring also in the pronotum (Fig. 139).

Meinert considers the elytra of Coleoptera to be the homologues of
the tegulæ of Lepidoptera and of Hymenoptera. He also calls
attention to the alula observed in Dyticus, situated at the base of
the elytra, but which is totally covered by the latter. The alulæ of
these beetles he regards as the homologues of the anterior wings of
Hymenoptera and Diptera. No details are given in support of these
views. (Ent. Tidskrift, i, 1880, p. 168.)

Hoffbauer (1892) also has suggested that the elytra are not the
homologues of the fore wings of other insects, but of the tegulæ.

Kolbe describes the alula of Dyticus as a delicate, membranous lobe
at the base of the elytra, but not visible when they are closed: its
fringed edge in Dyticus is bordered by a thickening forming a tube
which contains a fluid. The alula is united with the inner basal
portion and articulation of the wing-cover, forming a continuation
of them. Dufour considered that the humming noise made by these
beetles is produced by the alulets.

Hoffbauer finds no structural resemblances in the alulæ of Dyticus
to the elytra. He does not find “the least trace of veins.” They are
more like appendages of the elytra. Lacordaire considered that their
function is to prevent the disarticulation of the elytra, but
Hoffbauer thinks that they serve as contrivances to retain the air
which the beetle carries down with it under the surface, since he
almost always found a bubble of air concealed under it; besides,
their folded and fringed edge seems especially fitted for taking in
and retaining air. Hoffbauer then describes the tegulæ of the hornet
and finds them to be, not as Cholodkowsky states, hard, solid,
chitinous plates, but hollow. They are inserted immediately over the
base or insertion of the fore wings, being articulated by a
hinge-joint, the upper lamella extending into a cavity of the side
of the mesothorax, and connected by a hinge-like, articulating
membrane with the lower projection of the bag or cavity. The lower
lamella becomes thinner towards the place of insertion, is slightly
folded, and merges without any articulation into the thin, thoracic
wall at a point situated over the insertion of the fore wing. The
tegulæ also differ from the wings in having no muscles to move them,
the actual movements being of a passive nature, and due to the
upward and downward strokes of the wings.

Comstock adopts Meinert’s view that the elytra are not true fore
wings, but gives no reasons. (Manual, p. 495.)

Dr. Sharp,[25] however, after examining Dyticus and Cybister,
affirms that this structure is only a part of the elytron, to which
it is extensively attached, and that it corresponds with the angle
at the base of the wing seen in so many insects that fold their
front wings against the body. He does not think that the alula
affords any support to the view that the elytra of beetles
correspond with the tegulæ of Hymenoptera rather than with the fore
wings.

That the elytra are modified paraptera (tegulæ) is negatived by the
fact that the latter have no muscles, and that the elytra contain
tracheæ whose irregular arrangement may be part of the modified
degenerate structure of the elytra. Kolbe finds evidences of veins.
The question may also be settled by an examination of the structure
of the pupal wings. A study of a series of sections of both pairs of
wings of the pupa of Doryphora and of a Clytus convinces us that the
elytra are the homologues of the fore wings of other insects.

_e._ Development and mode of origin of the wings

=Embryonic development of the wings.=—The wings of insects are
essentially simple dorsal outgrowths of the integument, being
evaginations of the hypodermis. They begin to form in the embryo before
hatching, first appearing as folds, buds, or evaginations, of the
hypodermis, which lie in pouches, called peripodal cavities. They are
not visible externally until rather late in larval life, after the
insect, such as a grasshopper, has moulted twice or more times; while in
holometabolous insects they are not seen externally until the pupa state
is attained.

The subject of their origin is in a less satisfactory state than
desirable from the fact that at the outset the development of the wings
of the most generalized insects, such as Orthoptera, Termes, etc., was
not first examined, that of the most highly modified of any insects,
_i.e._ the Muscidæ, having actually been first studied.

In the course of his embryological studies on the Muscidæ (_Musca
comitoria_ and _Sarcophaga carnaria_) Weismann (1864) in examining the
larvæ of these flies just before pupation, found that the wings, as well
as the legs and mouth-appendages, developed from microscopic masses of
indifferent cells, which he called “imaginal discs.” From the six
imaginal discs or buds in the lower part of the thorax arise the legs,
while from four dorsal discs, two in the meso- and two in the
metathoracic segment, arise the fore and hind wings (Fig. 141.) These
imaginal buds, as we prefer to call these germs, usually appear at the
close of embryonic life, being found in freshly hatched larvæ.

[Illustration:

FIG. 140.—Imaginal buds in Musca,—_A_, in Corethra,—_B_, in
Melophagus,—_C_, in embryo of Melophagus; dorsal view of the head;
_b_, bud; _p_, peripodal membrane; _c_, cord; _hy_, hypodermis;
_cl_, cuticula; _st_, stomodæum; _v_, ventral cephalic, behind are
the two dorsal cephalic buds.—After Pratt.
]

As first observed by Weismann, the buds are, like those of the
appendages, simply attached to tracheæ and sometimes to nerves, in the
former case appearing as minute folds or swellings of the peritoneal
membrane of certain of the tracheæ. In Volucella the imaginal buds were,
however, found by Künckel d’Herculais to be in union with the
hypodermis. Dewitz detected a delicate thread-like stalk connecting the
peripodal membrane with the hypodermis, and Van Rees has since proved in
Musca, and Pratt in Melophagus, the connection of the imaginal buds with
the hypodermis (Fig. 140). These tracheal enlargements increase in size,
and become differentiated into a solid mass which corresponds to the
upper part of the mesothorax, while a tongue-shaped continuation becomes
the rudiment of the wing. During larval life the rudiments of the wings
crumple, thus forming a cavity. While the larva is transforming into the
pupa, the sheath or peripodal membranes of the rudimentary wings are
drawn back, the blood presses in, and thus the wings are everted out of
the peripodal cavities.

Due credit, however, should be given to Herold, as the pioneer in
these studies, who first described in his excellent work on the
development of _Pieris brassicæ_ (1815) the wing-germs in the
caterpillar after the third moult. This discovery has been
overlooked by recent writers, with the exception of Gonin, whose
statement of Herold’s views we have verified. Herold states that the
germs of the wings appear on the inside of the second and third
thoracic segments, and are recognized by their attachment to the
“protoplasmic network” (_schleimnetz_), which we take to be the
hypodermis, the net-like appearance of this structure being due to
the cell-walls of the elements of the hypodermal membrane. These
germs are, says Herold, also distinguished from the flakes of the
fat-body by their regular symmetrical form. Fine tracheæ are
attached to the wing-germs, in the same way as to the flakes of the
fat-body. It thus appears that Herold in a vague way attributes the
origin of these wing-germs, and also the germs of the leg, to the
hypodermis, since his schleimnetz is the membrane which builds up
the new skin. Herold also studied the later development of the
wings, and discovered the mode of origin of the veins, and in a
vague way traced the origin of the scales and hairs of the body, as
well as that of the colors of the butterfly.

Herold also says that as the caterpillar grows larger, and also the
wing-germs, “the larval skin in the region under which they lie
hidden is spotted and swollen,” and he adds in a footnote: “This is
the case with all smooth caterpillars marked with bright colors. In
dark and hairy caterpillars the swelling of the skin through the
growth of the underlying wing-germs is less distinct or not visible
at all” (pp. 29, 30).

It should be added that Malpighi, Swammerdam, and also Réaumur had
detected the rudiments of the wings in the caterpillar just before
pupation under the old larval skin. Lyonet (1760) also describes and
figures the four wing-germs situated in the second and third
thoracic segments, but was uncertain as to their nature. Each of
these masses, he says, is “situated in the fatty body without being
united to it, and is attached to the skin in a deep fold which it
makes there.” He could throw no certain light on their nature, but
says: “their number and situation leads to the supposition that they
may be the rudiments of the wings of the moth” (pp. 449, 450).

During the transformation into the pupa the imaginal buds unite and grow
out or extend along their edges, while the enveloping membrane
disappears. The rudimentary wings are now like little sacs, and soon
show a fusion of the two wing-membranes or laminæ with the veins, while
the tracheæ disappear, the places occupied by the tracheæ becoming the
veins. “Very early, as soon as the scales are indicated, begin in a very
peculiar way the fusion of the wing-laminæ. There occur openings in the
hypodermis into which the cells extend longitudinally and then laterally
give way to each other. Hence no complete opening is found, but the
epithelium appears by sections through a straight line sharply bordered
along the wingcavity. It is a continuous membrane formed of plasma which
I will call the ground membrane of the epithelium. Through this ground
membrane pass blood-corpuscles as well as blood-lymph.” (Schaeffer.)

[Illustration:

FIG. 141.—Anterior part of young larva of _Simulium sericea_, showing
the thoracic imaginal buds: _p_, prothoracic bud (only one not
embryonic); _w_, _w′_, fore and hind wing-buds; _l_, _l′_, _l″_,
leg-buds; _n_, nervous system; _br_, brain; _e_, eye; _sd_, salivary
duct; _p_, prothoracic foot.—After Weismann.
]

Afterwards (1866) Weismann studied the development of the wings in
_Corethra plumicornis_, which is a much more primitive and generalized
form than Musca, and in which the process of development of the wings is
much simpler, and, as since discovered, more as in other holometabolous
insects. He also examined those of Simulium (Fig. 141).

In Corethra, after the fourth and last larval moulting, there arises
at first by evagination and afterwards by invagination a cup-shaped
depression on each side in the upper part of the mesothoracic
segment within which the rudiment of the wings lies like a plug. The
wings without other change simply increase in size until, in the
transformation into the pupa by the withdrawal of the hypodermis,
the wings project out and become filled with blood, the tracheæ now
being wholly wanting, and other tissues being sparingly present.

[Illustration:

FIG. 142.—Section through thorax of a Tineid larva on sycamore,
passing through the 1st pair of wings (_w_): _ht_, heart; _i_,
œsophagus; _s_, salivary gland: _ut_, urinary tube; _nc_, nervous
cord; _m_, recti muscles; a part of the fat body overlies the heart.
_A_, right wing-germ enlarged.
]

These observations on two widely separate groups of Diptera were
confirmed by Landois, and afterwards by Pancritius, for the
Lepidoptera, by Ganin for the Hymenoptera, by Dewitz for Hymenoptera
(ants) and Trichoptera; also for the Neuroptera by Pancritius. In
the ant-lion (_Myrmeleon formicarius_) Pancritius found no rudiments
of the wings in larvæ a year old, but they were detected in the
second year of larval life, and do not differ much histologically or
in shape from those of Lepidoptera. In the Coleoptera and
Hymenoptera the imaginal buds appear rather late in larval life, yet
their structure is like that of Lepidoptera. In Cimbex the rudiments
of the wings are not found in the young larva, but are seen in the
semipupa, which stage lasts over six weeks.

[Illustration:

FIG. 143.—Section of the same specimen as in Fig. 142, but cut through
the second pair of wings (_w_): _i_, mid-intestine; _h_, heart;
_fb_, fat-body; _l_, leg; _n_, nervous cord.
]

The general relation of the rudiments (imaginal buds) of the wings of a
tineid moth to the rest of the body near the end of larval life may be
seen in Figs. 142, 143 (Tinea?), the sections not, however, showing
their connection with the hypodermis, which has been torn away during
the process of cutting. That the wing is but a fold of the hypodermis is
well seen in Fig. 144, of Datana, which represents a much later stage of
development than in Figs. 142 and 143, the larva just entering on the
semipupa stage.

In caterpillars of stage I, 3 to 4 mm. in length, Gonin found the
wing-germs as in Fig. 145, _A_ being a thickening of the hypodermis,
with the embryonic cells, _i.e._ of Verson, on the convex border. The
two leaves, or sides of the wing, begin to differentiate in stage II
(_C_, _D_), and in stage III the envelope is formed (_E_), while the
tracheæ begin to proliferate, and the capillary tracheæ or tracheoles at
this time arise (Fig. 145, _tc_). The wall of the principal trachea
appears to be resolved into filaments, and all the secondary branches
assume the appearance of bundles of twine. Landois regarded them as the
product of a transformation of the nuclei, but Gonin thinks they arise
from the entire cells, stating that from each cell arises a ball
(peloton) of small twisted tubes.

[Illustration:

FIG. 144.—Section through mesothoracic segment of _Datana ministra_,
passing through the wings (_w_): _c_, cuticula; _hyp_, hypodermis:
_ap_, apodeme; _dm_, dorsal longitudinal.—_vm_, ventral
longitudinal. muscles; _dmt_, depressor muscle of tergum; _t_,
trachea; _n_, nerve cords; _i_, intestine; _u_, urinary tubes; _l_,
insertion of legs.
]

As the large branches penetrate into the wing, the balls (pelotons) of
fine tracheal threads tend to unroll, and each of the new ramifications
of the secondary tracheal system is accompanied in its course by a
bundle of capillary tubes. This secondary system of wing-tracheæ, then,
arises from the mother trachea at the end of the third stage, when we
find already formed the chitinous tunic, which will persist through the
fourth stage up to pupation. It differs from the tracheoles in not
communicating with the air-passage; it possesses no spiral membrane at
the origin, and takes no part in respiration.

Gonin thus sums up the nature of the two tracheal systems in the
rudimentary wing, which he calls the provisional and permanent systems.
“The first, appearing in the second stage of the larva, comprises all
the capillary tubes, and arising from numerous branches passes off from
the lateral trunk of the thorax before reaching the wing; the second is
formed a little later by the direct ramification of the principal
branch.

“These two systems are absolutely independent of each other within the
wing. Their existence is simultaneous but not conjoint. One is
functionally active after the third moult; the other waits the final
transformation before becoming active.”

[Illustration:

FIG. 145.—_A_, section of wing-bud of larva of _Pieris brassicæ_ of
stage I, in front of the invagination pit. _B_, section passing
through the invagination pit. _C_, section of same in stage II,
through the invagination pit;—_D_, behind it, making the bud appear
independent of the thoracic wall. _E_, wing-bud at the beginning of
the 3d larval stage, section passing almost through the pedicel or
hypodermic insertion, the traces of which appear at _hi_; _h_,
hypodermis; _t_ or _tr_, trachea; _i_, opening of invagination;
_ec_, embryonic cells; _l_, external layer or envelope; _in_,
internal wall of the wing; _ex_, external wall; _s_, cell of a
tactile hair; _tc_, capillary tubes; _c_, cavity of
invagination.—After Gonin.
]

=Evagination of the wing outside of the body.=—We have seen that the
alary germs arise as invaginations of the hypodermis; we will now, with
the aid of Gonin’s account, briefly describe, so far as is known, the
mode of evagination of the wings. During the fourth and last stage of
the caterpillar of Pieris, the wings grow very rapidly, and undergo
important changes.

Six or seven days after the last larval moult the chitinous wall is
formed, the wing remaining transparent. It grows rapidly and its lower
edge extends near the legs. It is now much crumpled on the edge, owing
to its rapid growth within the limits of its own segment. Partly from
being somewhat retracted, and partly owing to the irregularity of its
surface, the wing gradually separates from its envelope, and the cavity
of invagination (Fig. 145, _c_) becomes more like a distinct or real
space. The outer opening of the alary sac enlarges quite plainly, though
without reaching the level of the edge of the wing.

This condition of things does not still exactly explain how the wing
passes to the outside of the body. Gonin compares these conditions to
those exhibited by a series of sections of the larva, made forty-eight
hours later, on a caterpillar which had just spun its girdle of silk. At
this time the wings have become entirely external, but, says Gonin, we
do not see the why or the how. The partition of the sac has disappeared,
and with it the cavity and the leaf of the envelope.

It appears probable that the partition has been destroyed, because
the space between the two teguments is strewn with numerous bits,
many of which adhere to the chitinous integument, while others are
scattered along the edges of the wings, in their folds, or between
the wings and the wall of the thorax.

Another series of sections showed that the exit of the fore wings
had been accomplished, while the hinder pair was undergoing the
process of eversion. In this case the partition showed signs of
degeneration: deformation of the nuclei, indistinct cellular limits,
pigmentation, granular leucocytes, and fatty globules.

After the destruction of the partition, what remains of the layer of
the envelope is destined to make a part of the thoracic wall and
undergoes for this purpose a superficial desquamation. The layer of
flattened cells is removed and replaced by a firmer epithelium like
that covering the other regions. It is this renewed hypodermis which
conceals the wing within, serves to separate it from the cavity of
the body, and gives the illusion of a complete change in its
situation. Other changes occur, all forming a complete regeneration,
but which does not accord with the description of Van Rees for the
Muscidæ. Finally, Gonin concludes that the débris scattered about
the wing comes from the two layers of the partition of the sac, from
the flattened hypodermis of the renewed envelope, from the chitinous
cuticle of the wing, and from the inner surface of the chitinous
integument.

He thinks that the metamorphosis of Pieris is intermediate between
the two types of Corethra and of Musca, established by Weismann, as
follows:

=Corethra.=—The wing is formed in a simple depression of the
hypodermic wall. No destruction.

=Pieris.=—The rudiment is concealed in a sac attached to the
hypodermis by a short pedicel. Destruction of the partition and its
replacement by a part of the thoracic wall by means of the imaginal
epithelium.

=Musca.=—The pedicel is represented by a cord of variable length,
whose cavity may be obliterated (Van Rees). The imaginal hypodermis
is substituted for the larval hypodermis, which has completely
disappeared, either by desquamation (Viallanes), or by histolytic
resorption (Van Rees).

=Extension of the wing; drawing out of the tracheoles.=—When it is
disengaged from the cavity, the wing greatly elongates and the creases
on its surface are smoothed out; the blood penetrates between the two
walls, and the cellular fibres, before relaxed and sinuous, are now
firmly extended.

Of the two tracheal systems, the large branches are sinuous, and they
are rendered more distinct by the presence of a spiral membrane; but the
two tunics are not separated as in the other tracheæ of the thorax;
moreover, the mouth choked up with débris does not yet communicate with
that of the principal trunk. The bundles of tracheoles on their part
form straight lines, as if the folds of the organ had had no influence
on them. As they have remained bound together, apart from the chitinous
membrane of the tracheal trunk, they become drawn out with this
membrane, at the time of exuviation, _i.e._ of pupation, and are drawn
out of the neighboring spiracle.

[Illustration:

FIG. 146.—Full-grown larva of _Pieris brassicæ_, opened along the
dorsal line: _d_, digestive canal; _s_, silk-gland; _g_, brain; _st
I_, prothoracic stigma; _st IV_, 1st abdominal stigma; _a_, _a′_,
germs (buds) of fore and hind wings; _p_, bud of prothoracic
segment;—those of the third pair are concealed under the
silk-glands; _I–III_, thoracic rings.—After Gonin.
]

“This is a very curious phenomenon, which can be verified
experimentally: if we cut off the wing, while sparing the larval
integument around the thoracic spiracles, we preserve the two
tracheal systems; the same operation performed after complete
removal of the larval skin does not give the secondary tracheal
system.” (Gonin.) Deceived by the appearance of the tracheoles while
still undeveloped, Landois and Pancritius, who have not mentioned
the drawing out of the capillaries of the larva, affirm that they
are destroyed by resorption in the chrysalis.

“The study of the tracheæ is closely connected with that of the
veins (nervures). It is well to guard against the error of Verson,
who mistakes for these last the large tracheal branches of the wing.
This confusion is easily explained; it proves that Verson had, with
us, recognized that the secondary system is, in the larva, exempt
from all respiratory function. Landois thought that the pupal period
was the time of formation of the veins. It seems to me probable that
they are derived from the sheath of the peritracheal spaces.”
(Gonin, pp. 30–33.)

[Illustration:

FIG. 147.—Left anterior wing of a larva 3 days before pupation. The
posterior part is rolled up: _st_, prothoracic stigma; _tr. i._,
internal tracheal trunk; _tr. e._, _tr. e.′_, external tracheal
trunk; _p_, cavity of a thoracic leg, with the imaginal bud
_b_.—After Gonin.
]

The appearance of the wing-germs in the fully grown caterpillar, as
revealed by simple dissection, is shown at Fig. 146; Fig. 147 represents
a wing of a larva three days before pupation, with the germ of a
thoracic leg.

[Illustration:

FIG. 148.—Graber’s diagrams for explaining the origin and primary
invagination of the hypodermis to form the germs of the leg (_b_),
and wings (_f_, _A-C_), and afterwards their evagination _D_, so
that they lie on the outside of the body. _E_, stage _B_, showing
the hypodermal cavities (_f_) and stalks connecting the germs with
the hypodermis (_z_).—After Graber.
]

[Illustration:

FIG. 149.—Section lengthwise through the left wing of mature larva in
_Pieris rapæ_: _t_, trachea; _hyp_, hypodermis; _c_, cuticula.—After
Mayer.
]

A. G. Mayer has examined the late development of the wings in _Pieris
rapæ_. Fig. 149 represents a frontal section through the left wing of a
mature larva and shows the rudiment of the wing, lying in its hypodermal
pocket or peripodal cavity. How the trachea passes into the rudimentary
wing, and eventually becomes divided into the branches, around which the
main veins afterwards form, is seen in Figs. 144, 147, 159.

The histological condition of the wing at this time is represented by
Fig. 151, the spindle-like hypodermal cells forming the two walls being
separated by the ground-membrane of Semper.

“While in the pupa state,” says Mayer, “the wing-membrane is thrown into
a very regular series of closely compressed folds, a single scale being
inserted upon the crest of each fold. When the butterfly issues from the
chrysalis, these folds in the pupal wings flatten out, and it is this
flattening which causes the expansion of the wings.... It is evident
that the wings after emergence undergo a great stretching and
flattening. The mechanics of the operation appears to be as follows. The
hæmolymph, or blood, within the wings is under considerable pressure,
and this pressure would naturally tend to enlarge the freshly emerged
wing into a balloon-shaped bag; but the hypodermal fibres (_h_) hold the
upper and lower walls of the wing-membrane closely together, and so,
instead of becoming a swollen bag, the wing becomes a thin flat one. And
thus it is that the little thick corrugated sac-like wings of the
freshly emerged insect become the large, thin, flat wings of the
imago.... The area of the wing of the imago of _Danais plexippus_ is 8.6
times that of the pupa. Now, as the wing of the young pupa has about 60
times the area of the wing in the mature larva, it is evident that in
passing from the larval state to maturity the area of the wings
increases more than 500 times.”

[Illustration:

FIG. 150.—Diagrammatic reproduction of Fig. 149 showing the wing-germ
in its peripodal cavity (_p_): _h’drm_, hypodermis; _tr_, trachea;
_cta_, cuticula; _a_, anterior end.—After Mayer.
]

[Illustration:

FIG. 151.—Section of the wing-germ, the upper and lower sides
connected by spindle-like hypodermic cells (_h_), forming the rods
of the adult wing; _mbr_, ground-membrane of Semper.—After Mayer.
]

_f._ The primitive origin of the wings

Farther observations are needed to connect the mode of formation of the
wings in the holometabolous insects with the more primitive mode of
origin seen in the hemimetabolous orders, but the former mode is
evidently inherited from the latter. Pancritius remarks that the
development of the rudiments of the wing in a hypodermal cavity is in
the holometabolic insects to be regarded as a later inherited character,
the external conditions causing it being unknown.

Fritz Müller was the first to investigate the mode of development of the
wings of the hemimetabolic insects, examining the young nymphs of
Termites. He regards the wings as evaginations of the hypodermis, which
externally appear as thoracic scale-like projections, into which enter
rather late in nymphal life tracheæ which correspond to the veins which
afterward arise.

[Illustration:

FIG. 152.—Rudimentary wing of young nymph of Blatta, with the five
principal veins developed.
]

The primitive mode of origin of the wings may, therefore, be best
understood by observing the early stages of those insects, such as the
Orthoptera and Hemiptera, which have an incomplete metamorphosis. If the
student will examine the nymphs of any locust in their successive
stages, he will see that the wings arise as simple expansions downward
and backward of the lateral edges of the meso- and metanotum. In the
second nymphal stage this change begins to take place, but it does not
become marked until the succeeding stage, when the indications of veins
begin to appear, and the lobe-like expansion of the notum is plainly
enough a rudimentary wing.

Graber[26] thus describes the mode of development of the wings in the
nymph of the cockroach:

“If one is looking only at the exterior of the process, he will
perceive sooner or later on the sides of the meso- and metathorax
pouch-like sacs, which increase in extent with the dorsal integument
and at the same time are more and more separated from the body.
These wing-covers either keep the same position as in the
flat-bodied Blattidæ, or in insects with bodies more compressed the
first rudiments hang down over the sides of the thorax. As soon as
they have exceeded a certain length, these wing-covers are laid over
on the back. However, if we study the process of development of the
wings with a microscope, by means of sections made obliquely through
the thorax, the process appears still more simple. The chief force
of all evolution is and remains the power of growth in a definite
direction. In regard to the skin this growth is possible in insects
only in this way; namely, that the outer layer of cells is increased
by the folds which are forced into the superficial chitinous skin.
These folds naturally grow from one moult to another in proportion
to the multiplication of the cells, and are not smoothed out until
after the moulting, when the outer resistance is overcome.

[Illustration:

FIG. 153.—Partial metamorphosis of _Melanoplus femur-rubrum_,
showing the five nymph stages, and the gradual growth of the
wings, which are first visible externally in 3, 3_b_,
3_c_.—Emerton _del._
]

“As, however, the first wing-layers depend upon the wrinkling of the
general integument of the body through the increase in the upper
layer, the further growth of the wings depends in the later stages
upon the wrinkling of the epidermis of the wing-membrane even, which
fact we also observe under the microscope when the new wings drawn
forth from the old covers appear at first to be quite creased
together. These wing-like wrinkles in the skin are not empty
pouches, but contain tissues and organs within, which are connected
with the skin, as the fat of the body, the network of tracheæ,
muscles, etc. Alongside the tracheæ, running through the former
wing-pouches and accompanied by the nerves, there are canals through
which the blood flows in and out.

[Illustration:

FIG. 154.—Stages in the growth of the wings of the nymph of _Termes
flavipes_: _A_, young; _a_, a wing enlarged. _B_, older nymph;
_b_, fore wing; _n_, a vein. _C_, wings more advanced;—_D_,
mature.
]

[Illustration:

FIG. 155.—Wings of nymph of Psocus.
]

“After the last moult, however, when the supply of moisture is very
much reduced in the wing-pouches, which are contracted at the
bottom, their two layers become closely united, and afterward grow
into one single, solid wing-membrane.

“These thick-walled blood-tubes arising above and beneath the upper
and lower membrane of the wing are the veins of the wings; the
development of the creased wings in the pupa of butterflies is
exactly like that of cockroaches and bugs. The difference is only
that the folds of integument furnishing the wings with an ample
store of material for their construction reach in a relatively
shorter time, that is the space of time between two moults, the same
extent that they would otherwise attain only in the course of
several periods of growth in the ametabolous insects.”

[Illustration:

FIG. 156.—Nymph of _Aphrophora permutata_, with enlarged view of the
wings and the veins: _pro_, pronotum; _sc_, mesoscutum; 1_ab_, 1st
abdominal segment.
]

Ignorant of Graber’s paper, we had arrived at the same result, after an
examination of the early nymph-stages of the cockroach, as well as the
locusts, Termites, and various Hemiptera. In all these forms it is
plainly to be seen that the wings are simply expansions, either
horizontal or partly vertical (where, as in locusts, etc., the body is
compressed, and the meso- and metanota are rounded downwards), of the
hinder and outer edge of the meso- and metanotum. As will be seen by
reference to the accompanying figures, the wings are notal (tergal)
outgrowths from the dorsal arch of the two hinder segments of the
thorax. At first, as seen in the young pupal cockroach (Fig. 152) and
locust (Fig. 153, also Figs. 154 and 156) the rudiments of the wings are
continuous with the notum. Late in nymphal life a suture and a
hinge-joint appear at the base of the wing, and thus there is some
movement of the wing upon the notum; finally, the tracheæ are well
developed in the wings, and numerous small sclerites are differentiated
at the base of the wing, to which the special muscles of flight are
attached, and thus the wings, after the last nymphal moult, have the
power of flapping, and of sustaining the insect in the air; they thus
become true organs of flight.

It is to be observed, then, that the wings in all hemimetabolous insects
are outgrowths from the notum, and not from the flanks or pleurum of the
thorax. There is, then, no structure in any other part of the body with
which they are homologous.

[Illustration:

FIG. 157.—Development of wings of Trichoptera: _A_, portion of
body-wall of young larva of Trichostegia; _ch_, cuticula, forming at
_r_ a projection into the hypodermis, _m_; _r_, and _d_, forming
thus the first rudiment of the wing. _B_, the parts in a larva of
nearly full size; _a_, _c_, _d_, _b_, the well-developed hypodermis
of the wing-germ separated into two parts by _r_, the penetrating
extension of the cuticula; _v_, mesoderm, _C_, wing-pad of another
Phryganeid freed from its case at its change to the pupa: _b_, _d_,
outer layer of the hypodermis (_m_) of the body-wall; _v_, inner
layer within nuclei.—After Dewitz, from Sharp.
]

The same may be said of the true Neuroptera, Trichoptera (Fig. 157), the
Coleoptera, and the Diptera, Lepidoptera, and Hymenoptera. As we have
observed in the house fly,[27] the wings are evidently outgrowths of the
meso- and metanotum; we have also observed this to be most probably the
case in the Lepidoptera, from observations on a Tortrix in different
stages of metamorphosis. It is also the case with the Hymenoptera, as we
have observed in bees and wasps;[28] and in these forms, and probably
all Hymenoptera, the wings are outgrowths of the scutal region of the
notum.

With these facts before us we may speculate as to the probable origin of
the wings of insects. The views held by some are those of Gegenbaur,
also adopted by Lubbock, and originally by myself.[29] According to
Gegenbaur:

“The wings must be regarded as homologous with the lamellar tracheal
gills, for they do not only agree with them in origin, but also in
their connection with the body, and in structure. In being limited
to the second and third thoracic segments they point to a reduction
in the number of the tracheal gills. It is quite clear that we must
suppose that the wings did not arise as such, but were developed
from organs which had another function, such as the tracheal gills;
I mean to say that such a supposition is necessary, for we cannot
imagine that the wings functioned as such in the lower stages of
their development, and that they could have been developed by having
such a function.”

[Illustration:

FIG. 158.—Changes in external form of the young larva of _Calotermes
rugosus_, showing, in _A_ and _B_, the mode of origin of the
wing-pads: _A_, newly hatched, with 9 antennal joints, × 8. _B_,
older larva, with 10 joints, × 8. _C_, next stage, with 11 joints, ×
8. _D_, larva, with twelve joints; the position of the parts of the
alimentary canal are shown: _v_, crop; _m_, stomach; _b_, “paunch”;
_e_, intestine; _r_, heart, × 16⁄3.—After Fritz Müller, from Sharp.
]

If we examine the tracheal gills of the smaller dragon-fly (Agrion), or
the May-flies, or Sialidæ, or Perlidæ, or Phryganeidæ, we see that they
are developed in a very arbitrary way, either at the end of the abdomen,
or on the sternum, or from the pleurum; moreover, in structure they
invariably have but a single trachea, from which minute twigs branch
out;[30] in the wings there are five or six main tracheæ, which give
rise to the veins. Thus, in themselves, irrespective of their position,
they are not the homologues of the gills. The latter are only developed
in the aquatic representatives of the Neuroptera and Pseudoneuroptera,
and are evidently adaptive, secondary, temporary organs, and are in no
sense ancestral, primitive structures from which the wings were
developed. There is no good reason to suppose that the aquatic Odonata
or Ephemerids or Neuroptera were not descendants of terrestrial forms.

To these results we had arrived by a review of the above-mentioned
facts, before meeting with Fritz Müller’s opinions, derived from a study
of the development of the wings of Calotermes (Fig. 158). Müller[31]
states that “(1) The wings of insects have not originated from ‘tracheal
gills.’ The wing-shaped continuations of the youngest larvæ are in fact
the only parts in which air tubes are completely wanting, while tracheæ
are richly developed in all other parts of the body.[32] (2) The wings
of insects have arisen from lateral continuations of the dorsal plates
of the body-segments with which they are connected.”

Now, speculating on the primary origin of wings, we need not suppose
that they originated in any aquatic form, but in some ancestral land
insect related to existing cockroaches and Termes. We may imagine
that the tergites (or notum) of the two hinder segments of the
thorax grew out laterally in some leaping and running insect; that
the expansion became of use in aiding to support the body in its
longer leaps, somewhat as the lateral expansions of the body aid the
flying squirrel or certain lizards in supporting the body during
their leaps. By natural selection these structures would be
transmitted in an improved condition until they became flexible,
_i.e._ attached by a rude hinge-joint to the tergal plates of the
meso- and metathorax. Then by continued use and attempts at flight
they would grow larger, until they would become permanent organs,
though still rudimentary, as in many existing Orthoptera, such as
certain Blattariæ and Pezotettix. By this time a fold or hinge
having been established, small chitinous pieces enclosed in membrane
would appear, until we should have a hinge flexible enough to allow
the wing to be folded on the back, and also to have a flapping
motion. A stray tracheal twig would naturally press or grow into the
base of the new structure. After the trachea running towards the
base of the wing had begun to send off branches into the rudimentary
structure, the number and direction of the future veins would become
determined on simple mechanical principles. The rudimentary
structures beating the air would need to be strengthened on the
front or costal edge. Here, then, would be developed the larger
number of main veins, two or three close together, and parallel.
These would be the costal, subcostal, and median veins. They would
throw out branches to strengthen the costal edge, while the branches
sent out to the outer and hinder edges of the wings might be less
numerous and farther apart. The net-veined wings of Orthoptera and
Pseudoneuroptera, as compared with the wings of Hymenoptera, show
that the wings of net-veined insects were largely used for
respiration as well as for flight, while in beetles and bees the
leading function is flight, that of respiration being quite
subordinate. The blood would then supply the parts, and thus
respiration or aëration of the blood would be demanded. As soon as
such expansions would be of even slight use to the insect as
breathing organs, the question as to their permanency would be
settled. Organs so useful both for flight and aëration of the blood
would be still further developed, until they would become permanent
structures, genuine wings. They would thus be readily transmitted,
and being of more use in adult life during the season of
reproduction, they would be still further developed, and thus those
insects which could fly the best, _i.e._ which had the strongest
wings, would be most successful in the struggle for existence. Thus
also, not being so much needed in larval life before the
reproductive organs are developed, they would not be transmitted
except in a very rudimentary way, as perhaps masses of internal
indifferent cells (imaginal discs), to the larva, being the rather
destined to develop late in larval and in pupal life. Thus the
development of the wings and of the generative organs would go hand
in hand, and become organs of adult life.[33]

=The development and structure of the tracheæ and veins of the
wing.=—The so-called veins (“nervures”) originate from fine tracheal
twigs which pass into the imaginal discs. A single longitudinal trachea
grows down into the wing-germ (Fig. 147), this branch arising through
simple budding of the large body-trachea passing under the rudiment of
the wing.

[Illustration:

FIG. 159.—Germ of a hind wing detached from its insertion, and
examined in glycerine: _i_, pedicel of insertion to the hypodermis;
_tr_, trachea; _b_, semicircular pad; _e_, enveloping membrane; _c_,
bundle of capillary tracheoles; the large tracheæ of the wing not
visible; they follow the course of the bundles of tracheoles.—After
Gonin.
]

Gonin states that before the tracheæ reach the wing they divide into a
great number of capillary tubes united into bundles and often tangled.
This mass of tracheæ does not penetrate into the wing-germ by one of its
free ends, but spreading over about a third of the surface of the wing,
separates into a dozen bundles which spread out fan-like in the interior
of the wing. (Fig. 159). These ramifications, as seen under the
microscope, are very irregular; they form here and there knots and
anastomoses. They end abruptly in tufts at a little distance from the
edge of the wing. A raised semicircular ridge (_b_) surrounds the base
of the wing, and within this the capillaries are formed, while on the
other side they are covered by a cellular layer.

Landois, he says, noticed neither the pedicel of the insertion of
the wing (_i_) nor the ridge (_b_). Herold only states that the
tracheæ pass like roots into the wing. Landois believed that they
formed an integral part of it. Dewitz and Pancritius used sections
to determine their situation.

Fig. 160 will illustrate Landois’ views as to the origin of the tracheæ
and veins. _A_ represents the germ of a hind wing attached to a trachea;
_c_ the elongated cells, in which, as seen at _B_, _c_, a fine tangled
tracheal thread (_t_) appears, seen to be magnified at _C_. The cell
walls break down, and the threads become those which pass through the
centre of the veins.

[Illustration:

FIG. 160.—Origin of the wings and their veins.—After Landois.
]

[Illustration:

FIG. 161.-Section of the “rib” of a vein: _c_, cord; _b_, twig.—After
Schaeffer.
]

=The wing-rods.=—Semper discovered in transverse sections of the wings,
what he called _Flügelrippen_; one such rib accompanying the trachea in
each vein. He did not discover its origin, and his description of it is
said to be somewhat erroneous. Schaeffer has recently examined the
structure, remarking: “I have surely observed the connection of this
cellular tube with the tracheæ. It is found in the base of the wing
where the lumen of the tracheæ is much widened. I only describe the
fully formed rib (_rippe_). In a cross-section it forms a usually
cylindrical tube which is covered by a very thin chitinous intima which
bears delicate twigs (Fig. 161). These twigs are analogous to the
thickened ridge of the tracheal intima. I can see no connection between
the branches of the different twigs. Through the ribs (_rippen_) extend
a central cord (_c_) which shows in longitudinal section a clear
longitudinal streaking. Semper regarded it as a nerve. But the
connection of the tube with the trachea contradicts this view. I can
only regard the cord as a separation-product of the cells of the walls.”

[Illustration:

FIG. 162.—Parts of a vein of the cockroach, showing the nerve (_n_) by
the side of the trachea (_tr_); _c_, blood-corpuscles.—After
Moseley.
]

=Other histological elements.=—These are the blood-lymph, corpuscles,
blood-building masses, and nerves. Schaeffer states that in the immature
pupal wings we find besides the large tracheæ, which are more or less
branched, and in the wing-veins at a later period, blood-corpuscles
which are more or less gorged with nutritive material, and also the
“balls of granules” of Weismann, which are perhaps the “single fat-body
cells” detected by Semper. Schaeffer also states that into the
hypodermal fold of the rudiments of the wings pass peculiar formations
of the fat-body and tracheal system, and connected with the fat-body are
masses of small cells which by Schaeffer are regarded as blood-building
masses.

Fine nerves have also been detected within the veins, Moseley stating
that a nerve-fibre accompanies the trachea in all the larger veins in
the insects he has examined (Fig. 162), while it is present in
Melolontha, where the trachea is absent.

LITERATURE ON THE WINGS

=Jurine, L.= Nouvelle méthode de classer les Hyménoptères et les
Dipterès. Genève, 1807, 4º pp. 319, 14 Pls.

—— Observations sur les ailes des Hyménoptères. (Mém. acad. Turin,
1820, xxiv, pp. 177–214.)

=Latreille, P. A.= De la formation des ailes des Insectes. (Mém. sur
divers sujets de l’histoire naturelle des Insectes, etc. Paris,
1819. Fasc. 8.)

—— De quelques appendices particuliers du thorax de divers Insectes.
(Mém. du Mus. d’Hist. nat., 1821, vii, pp. 1–21, 354–363.)

=Chabrier, J.= Essai sur le vol des insectes. (Mém. du Mus. d’Hist.
nat., 1820, vi, pp. 410–476; 1821, vii, pp. 297–372; 1822, viii, pp.
47–99, 349–403.) Separate, pp. 328, 13 Pls.

=Burmeister, Hermann.= Handbuch der entomologie, i, 1832, pp. 96–106,
263–267, 494–505.

—— Untersuchungen über die Flügeltypen der Coleopteren. (Abhandl. d.
naturf. Ges. Halle, 1854, ii, pp. 125–140, 1 Taf.)

=Romand, B. E. de.= Tableau de l’aile supérieure des Hyménoptères, 1
Pl. Paris, 1839. (Revue Zool., ii, pp. 339; Bericht von Erichson für
1839, pp. 54–56.)

=Lefebure, A.= Communication verbale sur la ptérologie des
Lépidoptères. (Annal. Soc. Ent. France, 1842, i, pp. 5–35, 3 Pls.
Also Revue Zool. Paris, 1842, pp. 52–58, 1 Pl.)

=Deschamps, B.= Recherches microscopiques sur l’organisation des
élytres des Coléoptères. (Ann. sc. nat., sér. 3, iii, 1845, pp.
354–363.)

=Heer, Oswald.= Die Insektenfauna der Tertiärgebilde von Oeningen und
Radaboj., 1847, 1. Teil, pp. 75–94.

=Newman, E.= Memorandum on the wing-rays of insects. (Trans. Ent. Soc.
London, ser. 2, iii, 1855, pp. 225–231.)

=Westwood, J. O.= Notes on the wing-veins of insects. (Trans. Ent.
Soc. London, ser. 2, iv, 1857, pp. 60–64.)

=Loew, H.= Die Schwinger der Dipteren. (Berlin, Entom. Zeitschr.,
1858, pp. 225–230.)

=Saussure, H. de.= Études sur l’aile des Orthoptères. (Ann. scienc.
nat., 5 sér. x, p. 161.)

=Schiner, J. R.= Ueber das Flügelgeäder der Dipteren. (Verhdl. k. k.
Zool.-bot., Ges. Wien, 1864, pp. 193–200, 1 Taf.)

=Hagen, H. A.= Ueber rationelle Benennung des Geäders in den Flügeln
der Insekten. (Stettin. Ent. Zeitung, 1870, xxxi, pp. 316–320, 1
Taf.)

—— Kurze Bemerkungen über das Flügelgeäder der Insekten. (Wiener
Entom. Zeit., 1886, v, pp. 311, 312.)

=Plateau, F.= Qu’est-ce que l’aile d’un insecte? (Stett. Ent. Zeit.
Jahrg. 32, 1871, pp. 33–42, 1 Taf. Journal d. Zool., ii, 1873, pp.
126–137.)

=Moseley, H. N.= On the circulation in the wing of _Blatta orientalis_
and other insects, etc. (Quart. Journ. Micr. Sc. 1871, xi, pp.
389–395, 1 Pl.)

=Roger, Otto.= Das Flügelgeäder der Kafer. Erlangen, 1875, 90 p.

=Rade, E.= Die westfalischen Donacien und ihre nachsten Verwandten. 3
Taf. (Vierter Jahresber. d. Westfal Prov.-Vereins f. Wiss. u. Kunst,
1876, pp. 52–87; Flügel, pp. 61–68.)

=Katter, F.= Ueber Inseckten, speziell Schmetterlingsflügel. (Entom.
Nachr., iv, 1878, pp. 279–281, 293–298, 304–309, 321–323.)

=Hofmann, Georg v.= Ueber die morphologische Deutung der
Insektenflügel. (Jahresber. d. akad.-naturwiss. Vereins, Graz, v
Jahrg., 1879, pp. 63–68.)

=Kolbe, H. J.= Das Flügelgeäder der Psociden und seine systematische
Bedeutung. (Stettin. Entom. Zeitung, 1880, pp. 179–186, 1 Taf.)

—— Die Zwischenraume zwischen den Punktstreifen der
punktiertgestreiften Flügeldecken der Coleoptera als rudimentare
Rippen aufgefasst. (Jahresber. zool. Sektion d. Westfal. Prov.-Ver.
f. Wiss. u. Kunst. Münster, 1886, pp. 57–59, 1 Taf.)

=Lee, A. Bolles.= Les balanciers des Diptères, leurs organes
sensifères et leurs histologie. (Recueil Zool. Suisse, i, 1885, pp.
363–392, 1 Pl.)

=Poppius, Alfred.= Ueber das Flügelgeäder der finnischen
Dendrometriden. 1 Taf. (Berl. Entom. Zeitschr., 1888, pp. 17–28.)

=Comstock, J. H.= On the homologies of the wing-veins of insects.
(American Naturalist, xxi, 1887, pp. 932–934.)

=Brauer, F.= Ansichten über die paläozoischen Insekten und deren
Deutung. (Annal. d. k. k. naturhist. Mus. Wien, Bd. i, 1886, pp.
86–126, 2 Taf.)

=Brauer, F., und J. Redtenbacher.= Ein Beitrag zur Entwicklung des
Flügelgeäders der Insekten. (Zool. Anz. 1888, pp. 443–447.)

=Redtenbacher, J.= Vergleichende Studien über das Flügelgeäder der
Insekten. 12 Taf. (Annalen d. k. k. naturhist. Hofmuseums zu Wien,
1886, i, pp. 153–231.)

=Schoch, G.= Miscellanea entomologica. I. Das Geäder des
Insektenflügels; II. Prolegomena zur Fauna dipterorum Helvetiae,
Wissenschaftl. (Beilage z. Programm d. Kantonsschule Zurich, 1889,
4º, 40 p.)

=Bondsdorff, A.= von. Ueber die Ableitung der Skulpturverhältnisse bei
den Deckflügeln der Coleopteren. (Zool. Anz., 1890, xiii Jahrg., pp.
342–346.)

=Spuler, Arnold.= Zur Phylogenie und Ontogenie des Flügelgeäders der
Schmetterlinge. (Zeitschr. wissens. Zool., liii, 597–646, 2 Taf.,
1892.)

Also the writings of Adolph, Bugnion, Calvert, Comstock, Diez, Giraud,
Gonin, Graber, Kellogg, Packard, Pratt, Scudder, Walsh.

_g._ Mechanism of flight

=Marey’s views on the flight of insects.=—As we owe more to Marey than
to any one else for what exact knowledge we have of the theory of flight
of insects, the following account is condensed from his work entitled
“Movement.” The exceedingly complicated movements of the wings would
lead us, he says, to suppose that there exists in insects a very complex
set of muscles of flight, but in reality, he claims, there are only the
two elevator and depressor muscles of each wing.[34] And Marey says that
when we examine more closely the mechanical conditions of the flight of
insects, we see that an upward and downward motion given by the muscles
is sufficient to produce all these successive acts, so well coordinated
with each other; the resistance of the air effecting all the other
movements. He also refers to the experiments of Giraud which prove that
the insect needs for flight a rigid main-rib and a flexible membrane.

[Illustration:

FIG. 163.—The two upper lines are produced by the contacts of a
drone’s wing on a smoked cylinder. In the middle are recorded the
vibrations of a tuning-fork (250 vibrations per second) for
comparison with the frequency of the wing movements. Below are seen
the movements of the wing of a bee.—After Marey.
]

If we take off the wing of an insect, and holding it by the small joint
which connects it with the thorax, expose it to a current of air, we see
that the plane of the wing is inclined more and more as it is subjected
to a more powerful impulse of the wind. The anterior nervure resists,
but the membranous portion which is prolonged behind bends on account of
its greater pliancy.

The wings of insects may be regarded simply as vibrating wires, and
hence the frequency of their movements can be calculated by the note
produced. Their movements can be recorded directly on a revolving
cylinder, previously blackened with smoke, the slightest touch of the
tip of the wing removing the black and exposing the white paper beneath;
Fig. 163 was obtained in this way. By this method it was calculated that
in the common fly the wings made 330 strokes per second, the bee 190,
the Macroglossus 72, the dragon-fly 28, and the butterfly (_Pieris
rapæ_) 9. Thus the smaller the species, the more rapid are the movements
of the wings.

[Illustration:

FIG. 164.—Appearance of a wasp flying in the sun: the extremity of the
wing is gilded.—After Marey.
]

The path or trajectory made by the tip of the wing is like a figure 8.
Marey obtained this by fastening a spangle of gold-leaf to the extremity
of a wasp’s wing. The insect was then seized with a pair of forceps and
held in the sun in front of a dark background, the luminous trajectory
shaping itself in the form of a lemniscate (Fig. 164).

To determine with accuracy the direction taken by the wing at
different stages of the trajectory, a small piece of capillary glass
tubing was blackened in the smoke of a candle, so that the slightest
touch on the glass was sufficient to remove the black coating and
show the direction of movement in each limb of the lemniscate. This
experiment was arranged as shown in Fig. 165. Different points on
the path of movement were tested by the smoked rod, and from the
track along which the black had been removed the direction of
movement was deduced. This direction is represented in the figure by
means of arrows.

[Illustration:

FIG. 165.—Experiment to test the direction of movement of an insect’s
wing: _a_, _a′_, _b_, _b′_, different positions of the smoked rod.
]

=Theory of insect flight.=—“The theory of insect flight,” says Marey,
“may be completely explained from the preceding experiments. The wing,
in its to-and-fro movement, is bent in various directions by the
resistance of the air. Its action is always that of an inclined plane
striking against a fluid and utilizing that part of the resistance which
is favorable to its onward progression.

“This mechanism is the same as that of a waterman’s scull, which as it
moves backwards and forwards is obliquely inclined in opposite
directions, each time communicating an impulse to the boat.”

The mechanism in the case of the insect’s wing is far simpler, however,
than in the process of sculling, since “the flexible membrane which
constitutes the anterior part of the wing presents a rigid border, which
enables the wing to incline itself at the most favorable angle.”

“The muscles only maintain the to-and-fro movement, the resistance of
the air does the rest, namely, effects those changes in surface
obliquity which determine the formation of an 8–shaped trajectory by the
extremity of the wing.”

[Illustration:

FIG. 166.—Bee flying about in the chamber of the apparatus.—After
Marey.
]

Lendenfeld has applied photography to determine the position of the
wings of a dragon-fly, and Marey has carried chronophotography
farther to indicate the normal trajectory of the wing, and to show
the position in flight. Fig. 166 shows a bee in various phases of
flight. “The insect sometimes assumes almost a horizontal position,
in which case the lower part of its body is much nearer the
object-glass than is its head, and yet both extremities are equally
well defined in the photograph. The successive images are separated
by an interval of 1⁄20 of a second (a long time when compared to the
total time occupied by a complete wing movement, _i.e._ 1⁄190 of a
second). And hence it is useless to attempt to gain a knowledge of
the successive phases of movement by examining the successive
photographs of a consecutive series representing an insect in
flight. Nevertheless an examination of isolated images affords
information of extreme interest with regard to the mechanism of
flight.

“We have seen that owing to the resistance of the air the expanse of
wing is distorted in various directions by atmospheric resistance.
Now, as the oscillations during flight are executed in a horizontal
plane, the obliquity of the wing-surface ought to diminish the
apparent breadth of the wing. This appearance can be seen in Fig.
167. There is here a comparison between two Tipulæ: the one in the
act of flight, the other perfectly motionless and resting against
the glass window.

[Illustration:

FIG. 167.—Illustration to show two Tipulæ, one of them remaining
motionless on the glass, and the other moving its limbs in
different directions, and setting its body at various
inclinations: the illustration only represents a small part of a
long series.—After Marey.
]

“The motionless insect maintains its wings in a position of vertical
extension; the plane is therefore at right angles to the axis of the
object-glass. The breadth of the wing can be seen in its entirety;
the nervures can be counted, and the rounding off of the extremities
of the wings is perfectly obvious. On the other hand, the flying
insect moves its wings in a horizontal direction, and owing to the
resistance of the air the expanse of the wings is obliquely
disposed, and only the projection of its surface can be seen in the
photograph. This is why the extremity of the wings appears as if it
were pointed, while the other parts look much narrower than normal.
The extent of the obliquity can be measured from the apparent
alteration in width, for the projection of this plane with the
vertical is the sine of the angle. From this it may be gathered that
the right wing (Fig. 168, third image) was inclined at an angle of
about 50° with the vertical, say 40° with the horizontal. This
inclination necessarily varies at different points of the trajectory
and must augment with the rapidity of movement; the obliquity
reaching its maximum in those portions of the wings which move with
the greatest velocity, namely, towards the extremities. The result
is that the wing becomes twisted at certain periods of the
movement.” (See the fourth image in Fig. 168.) The position of the
balancers seems to vary according to that of the wings. (Marey’s
Movement, pp. 253–257.)

[Illustration:

FIG. 168.—Tipula in the act of flying, showing the various attitudes
of the wings and the position of the balancers.
]

=Graber’s views as to the mechanism of the wings, flight, etc.=—Although
in reality insects possess but four wings, nature, says Graber,
evidently endeavors to make them dipteral. This end is attained in a
twofold manner. In the butterflies, bees, and cicadas, the four wings
never act independently of each other, as two individual pairs, but they
are always joined to a single flying plate by means of peculiar hooks,
rows of claws, grooved clamps, and similar contrivances proceeding from
the modified edges of the wings; indeed, this connection is usually
carried so far that the hind wings are entirely taken in tow by the
front, and consequently possess a relatively weak mechanism of motion.
The other mode of wing reduction consists in the fact that one pair is
thrown entirely out of employment. We observe this for instance in bugs,
beetles, grasshoppers, etc.

In the meantime, then, we may not trust to appearances. As their
development indeed teaches us, the wings as well as the additional
members must be regarded as actual evaginations of the common
sockets of the body, and in order especially to refute the prevalent
opinion that these wing-membranes are void of sensation, it should
be remembered that Leydig has proved the existence, as well as one
can be convinced by experiment, of a nerve-end apparatus in certain
basal or radical veins of the wing-membrane, which is very extensive
and complicated, and therefore indicates the performance of an
important function, perhaps of a kind of balancing sense, and also
that these same insect wings, with their delicate membrane, are very
easily affected by different outside agents, as, for instance,
warmth, currents of air, etc.

Usually in their inactive or passive state the wings are held off
horizontally from the body during flight, and are laid upon the back
again when the insect alights; but an exception occurs in most
butterflies and Neuroptera, among which the wing-joint allows only one
movement round the oblique and long axis of the wings. From this cause,
too, the insects just mentioned can unfold their wings suddenly.

[Illustration:

FIG. 169.—Anterior part of a Cicada for demonstrating the mechanism of
the articulation of the fore wing: _a_, articular head; _b_,
articular pan, frog, or cotyla; _g_, elastic band; _c_, _d_, _e_,
system of elastic rods; _r__{1}, _r__{2}, 1st and 2d abdominal
segments. _HF_, hind wings.—After Graber.
]

The transition of the wings from the active to the resting condition
seems to be by way of a purely passive process, which, therefore,
usually gives no trouble to the insect. The wing being extended by the
tractive power of the muscles, flies back, when this ceases, to its
former or resting posture by means of its natural elasticity, like a
spiral spring disturbed from its balance. The structure of this spring
joint is very different, however.

It usually consists (Fig. 169) of two parts. The wing can move
itself up and down in a vertical plane by means of the forward
joint, and at the same time can rotate somewhat round its long axis,
because the chitinous part mentioned above is ground off after the
fashion of a mandrel.

The hinder joint, at a greater distance from the body, virtually
consists of a rounded piece (_a_) capitate towards the outside, and
of a prettily hollowed socket (_b_) formed by the union of the thick
ribs of the hind wings, which slides round the head joint when the
wings snap back upon the back. The mechanism which causes this
turning is, however, of a somewhat complicated nature. The most
instrumental part of it is the powerful elastic band (_g_) which is
stretched over from the hinder edge of the mesothorax (_R_{2}_)
towards that of the wings. This membrane is extended by the
expansion of the wings, and draws them towards the body as soon as
the contraction of the muscles relaxes. This closing band of the
wings is assisted by a leverage system consisting of three little
chitinous rods (_c_, _d_, _e_), which at its joining presses inwards
on the body on one side, and on the hinder edge and head-joint of
the wing on the other.

We must, however, lay great stress on a few more kinds of wing
support.

[Illustration:

FIG. 170.—Mesothoracic skeleton of a stag beetle: _schi_, scutellum,
on each side of which is the articulation of the fore wing (_V_),
consisting of two small styliform processes (_v_, _h_) of the base
of the wing; _za_, tooth which fits into the cavity of the
wing-lock (_gr_); _l_, edge of the right wing, passing into the
corresponding groove (_fa_) of the left; _Di_, diaphragm for the
attachment of the tergal muscle of the metasternum; _Di_{1}_ (not
explained by author); _Ka_, acetabulum of the coxa (_Hü_); _Se_,
chitinous process for the attachment of the coxal muscle; _Fe_,
femur; _Sch_, tibia; _B_{2}_, sternum.—After Graber.
]

The wing-cases of beetles at their return from flight are joined
together like the shells of a mussel on the inside as well as to the
wedge-shaped plate (Fig. 170, _schi_) between their bases. There is
even a kind of clasp at hand for this purpose. The base of the wing,
that is, bears a pair of tooth-like projections (_za_), which fit
into the corresponding hollows of the little plate.

The commissure arising from the joining of the inner edges is
characteristic. Usually the wings on both sides interlock by means
of a groove, as in stag-beetles, but sometimes even, as in Chlamys,
after the manner of two cog-wheels, so that we have here also an
imitation of the two most prevalent methods which the cabinet-maker
uses in joining boards together.

The act of folding the broad hind wings among beetles is not less
significant than the arrangement of the fore wing. If we forcibly
spread out the former in a beetle which has just been killed and
then leave it to its own resources again, we observe the following
result: According to its peculiar mode of joining, the costal vein
on the fore edge approaches the mid or discoidal vein of the basal
half as well as the distal half of the wing, whence arises a
longitudinal fold which curves in underneath. Then the distal half
snaps under like the blade of a pocket knife and lies on the plane
of the costal edge of the wing, while it also draws after it the
neighboring wing-area. The soft hinder-edge portion turns in
simultaneously when this wing-area remains fixed to the body while
the costal portion is moving towards the middle line of the body.

The wing-membranes of almost all insects have, moreover, the
capability of folding themselves somewhat, and this power of
extending or contracting the wing-membrane at will is of great
importance in flight.

Yes, but how is the folded wing spread out again? The fact may be
shown more simply and easily than one might suppose, and may be most
plainly demonstrated even to a larger public by making an artificial
wing exactly after the pattern of the natural one, in which bits of
whalebone may take the place of veins and a piece of india rubber
the membrane spread out between them. The reader will be patient
while we just explain to him the act of unfolding of the membranous
wing of the beetle. The actual impulse for this unfolding is due to
the flexor muscles which pull on, and at the same time somewhat
raise the vein on the costal edge. By this means the membranous fold
lying directly behind the costal vein is first spread out. But since
this fold is connected with the longitudinal fold of the distal end
of the wing which closes like a blade, the wing-area last mentioned
which is attached to the middle fold of the wing by the elastic
spring-like diagonal vein becomes stretched out. The hinder rayed
portion adjacent to the body is, on the other hand, simply drawn
along when the wing stands off from the body.

In order to properly grasp the mechanism of the insect wing we must
again examine its mode of articulation to the body somewhat more
accurately.

[Illustration:

FIG. 171.—Longitudinal section through a Tipula: _a_, mouth; _an_,
antenna; _k_{3}_, maxillary palpus; _ol_, labrum; _oG_, brain;
_uG_, subœsophageal ganglion; _BG_, thoracic ganglion; _schl_,
œsophagus; _mD_, digestive canal; _Ov_, ovary; _vF_, fore wing;
_sch_, halter; _lm_, longitudinal—_b-r_, lateral muscles.—After
Graber.
]

If we select the halteres of a garden gnat (Tipula) at the moment of
extension, we shall find them to be formed almost exactly after the
pattern of our oars, since the oblong oar-blade passes into a
longitudinal handle. The pedicel of the balancer is formed by the
thick longitudinal primary veins of the wing-membrane. This pedicel
(Fig. 171) is implanted in the side of the thorax in such a manner
that the wing may be compared to the top of a ninepin. One may
think, and on the whole it is actually the fact, that the stiff
pedicel of the wing is inserted in the thoracic wall, and that a
short portion of it (Fig. 172), projects into the cavity of the
thorax. It is true there is no actual hole to be found in the
thoracic wall, as the intermediate space between the base or pedicel
of the wing and the aperture in the thorax is lined with a thin
yielding membrane, on which the wing is suspended as on an
axle-tree. According to this, therefore, the insect wing, as well as
any other appendage of arthropods, acts as a lever with two arms.
The reader can then conjecture what may be the further mechanism of
the wing machine. We only need now two muscles diametrically opposed
to each other and seizing on the power arm of the wing, one of which
pulls down the short wing arm, thereby raising the oar, while the
other pulls up the power arm. And indeed the raising of the wing
follows in the manner indicated, since a muscle (_hi_) is attached
to the end of the wing-handle (_a_) which projects freely into the
breast cavity by the contraction of which the power arm is drawn
down.

[Illustration:

FIG. 172.—Scheme of the flying apparatus of an insect: _mnl_,
thoracic walls; _ab_, wings; _c_, pivot; _d_, point of insertion
of the depressor muscle of the wing (_kd_);—_a_, that of the
elevator of the wing (_ai_); _rs_, muscle for expanding,—_ml_, for
contracting, the walls of the thorax.—After Graber.
]

[Illustration:

FIG. 173.—Muscles of the fore wing of a dragon-fly (_an_, _ax_),
exposed by removing the thoracic walls: _h_{1}_, _h_{2}_,
elevators,—_s_{1}_-_s_{5}_, depressors, of the wings (_s_{1}_,
_s_{2}_, rotators).—After Graber.
]

On the other hand, we have been entirely mistaken in reference to
the mechanism which lowers the wings. The muscle concerned, that is
_kd_, is not at all the antagonist of the elevator muscle of the
wing, since it is placed close by this latter, but nearer to the
thoracic wall. But then, how does it come to be the counterpart of
its neighbor? In fact, the lever of the wing is situated in the
projecting piece alone. The extensor muscle of the wing does not
pull on the power arm, but on the resistant arm on the other side of
the fulcrum (_c_). The illustration shows, however, how such a case
is possible. The membrane of the joint fastening the wing-stalk to
the thorax is turned up outwards below the stalk like a pouch. The
tendon of the flexor of the wing passes through this pouch to its
point of attachment (_c_) lying on the other side of the fulcrum
(_d_). Thus it is very simply explained how two muscles which act in
the same direction can nevertheless have an entirely contrary
working power.

This is in a way the bare physical scheme of the flying machine by
the help of which we shall more easily become acquainted with its
further details.

Dragon-flies are unquestionably the most suitable objects for the
study of the muscles pulling directly on the wing itself. If the
lateral thoracic wall (Fig. 173) be removed or the thorax opened
lengthwise there appears a whole storehouse of muscular cords which
are spread out in an oblique direction between the base of the wing
and the side of the thoracic plate. There is first to be
ascertained, by the experiment of pulling the individual muscles in
the line with a pincers, which ones serve for the lifting and which
for the lowering of the wings. In dragon-flies the muscles are
arranged in two rows and in such a way that the flexors or
depressors (_s_, 1 bis) cling directly to the thoracic wall (compare
also the muscle _dk_ in Fig. 172 and _se_ in Fig. 174), while the
raiser or extensor (_h_ 1, to _h_ 2, Fig. 172, _hi_ and Fig. 174
_he_) lie farther in. The form of the wing-muscles is sometimes
cylindrical, sometimes like a prism, or even ribbon-like. However,
the contracted bundles of fibres do not come directly upon the
joint-process we have described, but pass over often indeed at a
very considerable distance from them, into peculiar chitinous
tendons. These have the form of a cap-like plate, often serrate on
the edge, which is prolonged into a thread, which should be
considered as the direct continuation of the base of the wings. The
wings, therefore, sink down into the thoracic cavity as if they were
a row of cords ending in handles where the strain of the muscles is
applied.

[Illustration:

FIG. 174.—Transverse section through the thorax of a locust
(Stenobothrus): _b_{1}_, leg; _h_, heart; _ga_, ventral cord;
_se_, depressor,—_he_, elevator, of the wing (_fl_); _b-r_,
lateral muscles which expand the thoracic walls;—_lm_,
longitudinal muscles which contract them; _shm_, _uhm_, muscles to
the legs; _bg_, apodemes.—After Graber.
]

[Illustration:

FIG. 175.—Inner view of a portion of the left side of body of
_Libellula depressa_, showing a part of the mechanism of flight,
viz., some of the chitinous ridges at base of the upper wing, and
some of the insertions of the tendons of muscles: _A_, line of
section through the base of the upper wing, the wing being
supposed to be directed backwards. _C_, upper portion of mechanism
of the lower wing; _b_, lever extending between the pieces
connected with the two wings.—After von Lendenfeld, from Sharp.
]

As may be seen in Fig. 173, the contractile section of several of
the muscles of the wing (_s_{5}_) is extraordinarily reduced, while
its thread-like tendon is proportionately longer. This gradation
being almost like that of the pipes of an organ in the length of the
wing-muscles, as may so easily be observed in the large
dragon-flies, plainly indicates that the strain of the individual
muscles is quite different in strength, since, as the phenomenon of
flight demands it, the different parts of the base of the wing
become respectively relaxed in very dissimilar measure.

We have thus far discussed only the elevator and depressor muscles.
Other groups (_s^1s^3_) are yet to be added, however, crossing under
the first at acute angles, which when pulling the wing sidewise,
bring about in union with the other muscles a screw-like turning of
the wings.

While in dragon-flies all the muscles which are principally
influential in moving the wing are directly attached to it, and thus
evidently assert their strength most advantageously, the case is
essentially different with all other insects. Here, as has already
been superficially mentioned above, the entire set of muscles
affecting the wing is analyzed into two parts of which the smaller
only is usually directly joined to the wings, while the movement is
indirectly influenced by the remainder (Graber).

In the dragon-fly the two wings are “brought into correlative action
by means of a lever of unusual length existing amongst the chitinous
pieces in the body wall at the base of the wings (Fig. 175, _b_).
The wing-muscles are large; according to von Lendenfeld there are
three elevator, five depressor, and one abductor muscles to each
wing. He describes the wing-movements as the results of the
correlative action of numerous muscles and ligaments, and of a great
number of chitinous pieces connected in a jointed manner” (Sharp).

If again we take the longitudinal section of the thoracic cavity of
gnats in Fig. 171, we shall perceive a compactly closed system of
muscular bars intersecting each other almost at right angles and
interlaced with a tangled mass of tracheæ, some of which muscles
extend (_lm_) longitudinally, that is from the front to the back,
while others (_b-r_) stretch out in a vertical direction, that is
between the plates of the abdomen and back.

In order that we may more easily comprehend this important muscular
apparatus we will illustrate the thoracic cavity of insects by an
elastic steel ring (Fig. 172), to which we may affix artificial
wings. If this ring be pressed together from above downward, along
the line _rs_, thus imitating the pulling of the vertical or lateral
thoracic muscles, then the wings on both sides spring up. This is to
be explained by the fact that through this manipulation a pressure
is exerted on the lifting power arm of the wings. If, on the other
hand, the ring be compressed on the sides (_ml_), which is the same
thing as if the longitudinal muscles contracted the thorax from
before backward, and thus arched it more, then the wings are
lowered.

Agrioninæ, according to Kolbe, can fly with the fore pair of wings
or with the hind pair almost as well as with both pairs together.
Also the wings of these insects can be cut off before the middle of
their length without injuring their power of flight. Butterflies,
Catocalæ, and Bombycidæ fly after the removal of the hind wings.
Also the balancers of the Diptera must be useful in flying, since
their removal lessens the power of flight.

Chabrier regarded the under sides of the shell-like extended
wing-covers of the beetles as wind-catchers, which, seized by wind
currents, carry the insect through the air. We may also consider the
wing-covers as regulators of the centre of gravity of flight.

The observations of insects made by Poujade (Ann. Soc. Ent., France,
1887, p. 197) during flight teaches us, says Kolbe, that in respect
to the movement during flight of both pairs of wings, they may be
divided into two categories:—

1. Into those where both pairs of wings (together), either united,
and also when separated from each other, perform flight. Such are
the Libellulidæ, Perlidæ, Sialidæ, Hemerobiidæ, Mymeleonidæ,
Acridiidæ, Locustidæ, Blattidæ, Termitidæ, etc.

2. Into those whose fore and hind wings act together like one wing,
since they are connected by hooks (hamuli), as in certain
Hymenoptera, or are attached in other ways. Here belong Hymenoptera,
Lepidoptera, Trichoptera, Cicadidæ, Psocidæ, etc.

The musculature of the mesothorax and metathorax is similar in those
insects both of whose pairs of wings are like each other, and act
independently during flight, viz. in the Libellulidæ. On the other
hand, in the second category, where the fore and hind wings act as a
single pair and the fore wings are mostly larger than the hinder
(except in most of the Trichoptera), the musculature of the
mesothorax is more developed than that of the metathorax.

To neither category belong the beetles, whose wing-covers are
peculiar organs of flight, and not for direct use, and the Diptera,
which possess but a single pair of wings. In the beetles the hind
wings, in the Diptera the fore wings, serve especially as organs of
flight. It may be observed that the Diptera are the best fliers, and
that those insects which use both pairs of wings as a single pair
fly better than those insects whose two pairs of wings work
independently of each other. An exception are the swift-flying
Libellulidæ, whose specially formed muscles of flight explain their
unusual capabilities for flying (Kolbe).

LITERATURE ON FLIGHT

=Marey, E. J.= La machine animale. Locomotion terrestre et aërienne.
Paris, 1874.

—— Mémoire sur le vol des insectes et des oiseaux. (Annal. Scienc.
natur., 5 sér., Zool. xii, 1869, pp. 49–150; 5 sér., Zool. xv, 1872,
42 Figs.)

—— Note sur le vol des insectes. (Compt. rend. et Mém. Soc. d. Biol.
Paris, 4 sér., v, 1869, C. R. pp. 136–139.)

—— Recherches sur le mécanisme du vol des insectes. (Journal de
l’Anatomie et de la Physiologie, 6 Année, 1869, pp. 19–36, 337–348.)

—— Animal mechanism. New York, 1879, pp. 180–209.

—— Movement. New York, 1895, pp. 239–274.

=Hartings.= Ueber den Flug. (Niederland. Archiv f. Zoologie, iv,
Leiden, 1877–78.)

=Lucy.= Le vol des oiseaux, chauvesouris et insectes. Paris.

=Tatin, V.= Expériences physiologiques et synthétiques sur le
mécanisme du vol. (Ecole prat. d. haut. étud. Physiol. expérim.
Trav. du laborat. de Marey, 1877, pp. 293–302.)

—— Expériences sur le vol mécanique. (Ibid., 1876, pp. 87–108.)

=Bellesme, Jousset de.= Recherches expérimentales sur les fonctions du
balancier chez les insectes Diptères, Paris, 1878, 96 pp., Figs.

—— Sur une fonction de direction dans le vol des insectes. (Compt.
rend., lxxxix, 1879, pp. 980–983.)

=Pettigrew, J. Bell.= On the mechanical appliances by which flight is
attained in the animal kingdom. (Trans. Linn. Soc., 1868, xxvi, Pt.
I, pp. 197–277, 4 Pl.)

—— On the physiology of wings. (Trans. Roy. Soc. Edinburgh, 1871,
xxvi, pp. 321–446.)

=Krarup-hansen, C. J. L.= Beitrag zu einer Theorie des Fluges der
Vogel, Insekten und Fledermause. (Copenhagen u. Leipzig, Fritsch,
1869, 48 pp.)

=Lendenfeld, R. V.= Der Flug der Libellen. (Sitzungsber. d. kais.
Akad. d. Wiss. Wien, lxxxiii, 1881, pp. 289–376, 7 Taf.; Zool. Anz.,
1880, p. 82.)

=Girard, M.= Note sur diverses expériences relatives à la fonction des
ailes chez les insectes. (Ann. Soc. Ent. France, 4 sér., ii, 1862,
pp. 154–162.)

=Mühlhäuser, F. A.= Ueber das Fliegen der Insekten. (22. bis 24.,
Jahresb. d. Pollichia, Dürkheim, 1866, pp. 37–42.)

=Plateau, Félix.= Recherches expérimentales sur la position du centre
de gravité chez les insectes. (Archiv d. Scienc. phys. et natur. d.
Genève, Nouv. période, xliii, 1872, pp. 5–37.)

=Plateau, Félix.= Ueber die Lage des Schwerpunktes bei den Insekten.
Auszug. (Naturforscher v. Sklarek, v. Jahrg., 1872, pp. 112–113.)

—— Recherches physico-chimiques sur les articulés aquatiques. (Bull.
d. l’Acad. Roy. Belg., xxxiv, 1872, pp. 1–50, Fig.)

—— Qu’est-ce que l’aile d’un insecte? (Stett. Ent. Zeit., 1871, pp.
33–42, Pl.)

—— L’aile des insectes. (Journ. d. Zool., ii, 1873, pp. 126–137.)

=Perez, J.= Sur les causes de bourdonnement chez les insectes.
(Comptes rend., lxxxvii, p. 535, Paris, 1878.)

=Strasser, Hans.= Mechanik des Fluges. (Archiv f. Anat. u. Phys.,
1878, p. 310–350, 1 Taf.)

—— Ueber die Grundbedingungen der aktiven Locomotion. (Abhandl. d.
naturf. Gesellsch., Halle, 1880, xv, pp. 121–196, Figs.)

=Moleyre, L.= Recherches sur les organes du vol chez les insectes de
l’ordre des Hemiptères. (Compt. rend. de l’Acad. d. Scienc. de
Paris, 1882, xcv, pp. 349–352.)

=Amans, P.= Essai sur le vol des insectes. (Revue d. Sc. Nat.
Montpellier, 3 sér., ii, 1883, pp. 469–490, 2 Pl.; iii, 1884, pp.
121–139, 3 Pl.)

—— Étude de l’organe du vol chez les Hyménoptères. (Ibid., iii, pp.
485–522, 2 Pl.)

—— Comparaisons des organes du vol dans la série animale. Des organes
du vol chez les insectes. (Annal. d. Scienc. nat. Zool., 6 sér.,
xix, pp. 1–222, 8 Pl.)

=Mullenhoff, K.= Die Grosse der Flugflächen. (Pflüger’s Archiv f. d.
ges. Physiologie, 1884, xxxv, pp. 407–453.)

—— Die Ortsbewegungen der Tiere. (Wissensch. Beil. z. Programm d.
Andreas-Realgymnas. Berlin, 1885, 19 pp.)

=Poujade, G. A.= Note sur les attitudes des insectes pendant le vol.
(Ann. Soc. Ent. France, 1884, 6 sér., iv, pp. 197–200, 1 Pl.)

=Krancher, O.= Die Töne der Flügelschwingungen unserer Honigbiene.
(Deutscher Bienenfreund, 1882, 18. Jahrg., pp. 197–204.)

=Landois, H.= Ueber das Flugvermögen der Insekten. (Natur. und
Offenbarung, vi, 1860, pp. 529–540.)

=Ungern-Sternberg, von.= Betrachtungen über die Gesetze des
Fluges. (Zeitschr. d. Deutschen Vereins z. Förderung d.
Luftschiffahrt-Naturwissensch. Wochenschrift v. Potonie, iv,
1889, p. 158.)

=Baudelot, E.= Du mecanisme suivant lequel s’effectue chez les
Coléoptères le retract des ailes inférieures sous les élytres au
moment du passage a l’état de repos. (Bull. Soc. d. Scienc. nat.,
Strasbourg, 1 Année, 1868, pp. 137–138.)

=Ris, Fr.= Die schweizerischen Libellen. Schaffhausen, 1885. (Beiheft
der Mitteil. d. Schweiz. Ent. Ges., vii, pp. 35–84.)

THE ABDOMEN AND ITS APPENDAGES

[Illustration:

FIG. 176.—Abdomen of _Termes flavipes_: 1–10, the ten tergites; 1–9,
the nine urites; _c_, cercopod.
]

[Illustration:

FIG. 177.—End of abdomen of _Panorpa debilis_ drawn out, the chitinous
pieces shaded: _L_, lateral, _D_, dorsal view; _c_, jointed
cercopoda.—Gissler _del._
]

In the abdomen the segments are more equally developed than elsewhere,
retaining the simple annular shape of embryonic life, and from their
generalized nature their number can be readily distinguished (Fig. 176).
The tergal and sternal pieces of each segment are of nearly the same
size, the tergal often overlapping the sternal (though in the Coleoptera
the sternites are larger than the tergites), while there are no pleural
pieces, the lateral region being membranous when visible and bearing the
stigmata (Fig. 177, _L_). In the terminal segments beyond the genital
outlet, however, there is a reduction in and loss of segments,
especially in the adults of the metabolous orders, notably the Panorpidæ
(Fig. 177), Diptera, and aculeate Hymenoptera; in the Chrysididæ only
three or four being usually visible, the distal segments being reduced
and telescoped inward.

The typical number of abdominal segments (uromeres), _i.e._ that
occurring in each order of insects, is ten; and in certain families of
Orthoptera, eleven. In the embryos, however, of the most generalized
winged orders, Orthoptera (Fig. 199), Dermaptera, and Odonata, eleven
can be seen, while Heymons has recently detected twelve in blattid and
Forficula embryos, and he claims that in the nymphs of certain Odonata
there are twelve segments, the twelfth being represented by the anal or
lateral plates. It thus appears that even in the embryo condition of the
more generalized winged insects, the number of uromeres is slightly
variable.

We have designated the abdomen as the _urosome_; the abdominal
segments of insects and other Arthropods as _uromeres_, and the
sternal sclerites as _urosternites_, farther condensed into
_urites_. (See Third Report U. S. Entomological Commission, 1883,
pp. 307, 324, 435, etc.)

[Illustration:

FIG. 178.—Nymph of the pear tree Psylla, with its glandular
hairs.—After Slingerland. Bull. Div. Ent. U. S. Dep. Agr.
]

The reduction takes place at the end of the abdomen, and is usually
correlated with the presence or absence of the ovipositor. In the more
generalized insects, as the cockroaches, the tenth segment is, in the
female, completely aborted, the ventral plate being atrophied, while the
dorsal plate is fused during embryonic life, as Cholodkowsky has shown,
with the ninth tergite, thus forming the suranal plate.

In the advanced nymph of Psylla the hinder segments of the abdomen
appear to be fused together, the traces of segmentation being
obliterated, though the segments are free in the first stage and in
the imago (Fig. 178). It thus recalls the abdomen of spiders, of
Limulus, and the pygidium of trilobites.

=The median segment.=—There has been in the past much discussion as to
the nature of the first abdominal segment, which, in those Hymenoptera
exclusive of the phytophagous families, forms a part of the thorax, so
that the latter in reality consists of four segments, what appearing to
be the first abdominal segment being in reality the second.

Latreille and also Audouin considered it as the basal segment of the
abdomen, the former calling it the “segment médiaire,” while Newman
termed it the “propodeum.” This view was afterward held by Newport,
Schiödte, Reinhard, and by the writer, as well as Osten-Sacken,
Brauer, and others. The first author to attempt to prove this by a
study of the transformations was Newport in 1839 (article
“Insecta”). He states that while the body of the larva is in general
composed of thirteen distinct segments, counting the head as the
first, “the second, third, fourth, and, as we shall hereafter see,
in part also the fifth, together form the thorax of the future
imago” (p. 870). Although at first inclined to Audouin’s opinion, he
does not appear to fully accept it, yet farther on (p. 921) he
concludes that in the Hymenoptera the “fifth” segment (first
abdominal) is not in reality a part of the true thorax, “but is
sometimes connected more or less with that region, or with the
abdomen, being intermediate between the two. Hence we have ventured
to designate it the _thoracico-abdominal_ segment.” Had he
considered the higher Hymenoptera alone, he would undoubtedly have
adopted Latreille’s view, but he saw that in the saw-flies and
Lepidoptera the first abdominal segment is not entirely united with
the thorax, being still connected with the abdomen as well as the
thorax. Reinhard in 1865 reaffirmed Latreille’s view. In 1866 we
stated from observations on the larvæ made three years earlier, that
during the semipupa stage of Bombus the entire first abdominal
segment is “transferred from the abdomen to the thorax with which it
is intimately united in the Hymenoptera,” and we added that we
deemed this to be “the most essential zoölogical character
separating the Hymenoptera from all other insects.” (See Fig. 93,
showing the gradual transfer and fusion of this segment with the
thorax.) In the saw-flies the fusion is incomplete, as also in the
Lepidoptera, while in the Diptera and all other orders the thorax
consists of but three segments. (See also pp. 90–92.)

[Illustration:

FIG. 179.—Abdomen of _Machilis maritima_, ♀, seen from beneath: the
left half of the 8th ventral plate removed; I-IX, abdominal
segments; _c_, cercopoda; _cb_, coxal glands; _hs_, coxal stylets;
_lr_, ovipositor.—After Oudemans, from Lang.
]

=The cercopoda.=—We have applied this name to the pair of anal cerci
appended to the tenth abdominal segment, and which are generally
regarded as true abdominal legs. As is now well known, the embryos of
insects of different orders have numerous temporary pairs of abdominal
appendages which arise in the same manner, have the same embryonic
structure, and are placed in a position homologous with those of the
thorax. In the embryo of Œcanthus rudimentary legs appear, as shown by
Ayers, on the first to tenth abdominal segment, the last or tenth pair
becoming the cercopoda; and similar rudimentary appendages have been
detected in the embryos of Coleoptera, Lepidoptera, and Hymenoptera
(Apidæ). Cholodkowsky has observed eleven pairs of abdominal appendages
in Phyllodromia.

They are very long and multiarticulate in the Thysanura (Fig. 179). In
the Dermaptera they are not jointed and are forcep-like. It should also
be observed that in the larva or Sisyra (Fig. 181) there are seven pairs
of 5–jointed abdominal appendages, though these may be secondary
structures or tracheal gills. In the Perlidæ and the Plectoptera
(Ephemeridæ), they are very long, sometimes over twice as long as the
body, and composed of upward of 55 joints; they also occur in the
Panorpidæ (Fig. 177). In the dragon-flies the cerci are large, but not
articulated, and serve as claspers or are leaf-like[35] (Fig. 180). In a
few Coleoptera, as the palm-weevil (_Rhynchophorus phœnicis_), Cerambyx,
Drilus, etc., the so-called ovipositor ends in a hairy, 1–jointed,
palpiform cercus. Short 25–jointed cercopoda are present in Termitidæ,
and 2–jointed ones in Embiidæ.

[Illustration:

FIG. 180.—End of abdomen of _Æschna heros_, ♀: _ur_, urosternite;
_or_, outer, _ir_, inner styles of the ovipositor; 11, 11th
abdominal segment; _c_, cercopod.
]

[Illustration:

FIG. 181.—Larva of Sisyra, from beneath. _B_, an abdominal
appendage.—After Westwood, from Sharp.
]

[Illustration:

FIG. 182.—Cercopoda (P) of Mantis.—After Lacaze-Duthiers.
]

The anal cerci are present in the Orthoptera and, when multiarticulate,
function as abdominal antennæ. They are longest in the Mantidæ (Fig.
182); they also occur in the larva of the saw-fly, Lyda (Fig. 183). Dr.
A. Dohrn has stated that the cerci of Gryllotalpa are true sensory
organs, and we have called those of the cockroach abdominal antennæ,
having detected about ninety sacs on the upper side of each joint of the
stylets, which are supposed to be olfactory in nature, and which are
larger and more numerous than similar sacs or pits in the antennæ of the
same insect.[36] From his experiments upon decapitated cockroaches,
Graber concluded that these cerci were organs of smell.

[Illustration:

FIG. 183.—Lyda larva: _a_, head; _b_, end of body seen from above;
_c_, from side, with cercopod.
]

Haase regarded these appendages, from their late development and
frequent reduction, as old inherited appendages which are
approaching atrophy through disuse.

Cholodkowsky states that Tridactylus, a form allied to Gryllotalpa,
bears on the tenth abdominal segment two pairs of cerci (ventral and
dorsal), and that the ventral pair may correspond to the atrophied
appendages of the tenth embryonic segment of Phyllodromia, with
which afterward the eleventh segment becomes fused.

The cercopods are not necessarily confined to the eleventh or to the
tenth segment, for when there are only nine segments, with the
vestige of a tenth, as in Xiphidium, they arise from the ninth
uromere, and in the more modern cockroaches, as Panesthia, in which
there are but seven entire segments, they are appended to the last
or eighth uromere.

[Illustration:

FIG. 184.—Anabrus, ♀, side-view, dissected; showing the relative size
of the ovipositor: _c_, the minute cercopod.—Kingsley _del._
]

As to the homology and continuity of these cercopods with the ventral
outgrowths of the embryo, several embryologists, notably Wheeler, are
emphatic in regarding them as such. It thus appears that either the
embryonic appendages of the seventh or eighth, ninth or tenth uromere
may persist, and form the cercopoda of the adult.

=The ovipositor.=—The end of the oviduct is guarded by three pairs of
chitinous, unjointed styles closely fitted together, forming a strong,
powerful apparatus for boring into the ground or into leaves, stems of
plants, the bodies of insects, or even into solid wood, so that the eggs
may be deposited in a place of safety. In the ants, wasps, and bees the
ovipositor also functions as a sting, which is further provided with a
poison-sac.

Morphologically, the ovipositor is composed of three pairs of unjointed
styles (_rhabdites_ of Lacaze-Duthiers, _gonapophyses_ of Huxley), which
are closely appressed to or sheathed within each other, the eggs passing
out from the end of the oviduct, which lies, as Dewitz states, between
the two styles of the lowest or innermost pair, and under the cross-bars
or at the base of the stylets mentioned; the styles or blades spreading
apart to allow of the passage of the egg.

[Illustration:

FIG. 185.—Saw of Hylotoma: _a_, lateral scale; _i_, saw; _f_, gorget;
7_t_, 7th tergite; 6_s_, 6th sternite; _ov_, oviduct; _in_,
intestine.—After Lacaze-Duthiers.
]

The ovipositor is best developed in the Thysanura (Fig. 179, Campodea
excepted), in Orthoptera (Fig. 184), in the Odonata, Hemiptera, certain
Physapoda, Rhaphiidæ, and in the phytophagous Hymenoptera, where it is
curiously modified to form a rather complicated saw for cutting slits in
wood or leaves (Fig. 185). It is wanting or quite imperfect in
Coleoptera, Diptera, and Lepidoptera.

Morphologically, the ovipositor appears to be formed out of the
abdominal appendages of the seventh, eighth, and ninth segments of the
female, which, instead of disappearing in the orders first mentioned,
persist as permanent styles.

Wheeler asserts from his study of the embryonic development of Xiphidium
“there can be no doubt concerning the direct continuity of the embryonic
appendages with the gonapophyses.” He goes on to say:—

“One embryo, which had just completed katatrepis, still showed
traces of all the abdominal appendages. The pairs on the eighth,
ninth, and tenth segments were somewhat enlarged. In immediately
succeeding stages the appendages of the second to sixth segments
disappear; the pair on the seventh disappear somewhat later. Up to
the time of hatching the gonapophyses could be continuously traced,
since in Xiphidium there is no flexure of the abdomen, as in other
forms, to obscure the ventral view of the terminal segments. From
the time of hatching Dewitz has traced the development of the
ovipositor in another locustid (_Locusta viridissima_), so that now
we have the complete history of the organ.”

Heymons, however, is inclined to believe that they are simply hypodermal
outgrowths.

[Illustration:

FIG. 186.—Ideal plan of the structure of the ovipositor to illustrate
Lacaze-Duthiers’ view: _b_, 8th tergite; _c_, epimerum; _a′_, _a_,
two pieces forming the outer pair of rhabdites; _i_, the 2d pair, or
stylets; and _f_, the inner pair, or sting; _d_, support of sting;
_e_, piece supporting the stylet; _R_, anus; _o_, outlet of oviduct.
The 7th, 8th, and 9th sternites are aborted.—After Lacaze-Duthiers.
]

The first to study the morphology of the ovipositor was Lacaze-Duthiers,
who referred their origin to the partially atrophied dorsal or ventral
sclerites of one of the last abdominal segments; a view accepted by
Gerstaecker[37] (Figs. 186, 187). The present writer (1866), however,
showed that the sting of Bombus was not formed of the reduced pieces of
the segments themselves, but arose from special outgrowths on the
ventral side of the eighth and ninth abdominal segments. These
appendages he did not at first regard as the homologues of the limbs,
until in 1871, after studying the origin of the spring of the Podurans
(Isotoma), he found that it was a true jointed appendage and therefore a
homologue of a pair of the styles forming the ovipositor of the winged
insects, and that the three pairs of styles of the latter were
homologues of the thoracic legs and cephalic appendages. The view was
stated in the Guide to the Study of Insects. (See also Amer. Nat.,
March, 1871, p. 6.) Kraepelin also affirms that the styles of the
ovipositor are segmental appendages and homologues of the antennæ, wings
(_sic_), and legs.

[Illustration:

FIG. 187.—1, abdomen of Cynips, showing the great dorsal segment, the
peduncle, and the position of the ovipositor within; 2, the entire
ovipositor; _a_, lateral scale; _a′_, its valve; _b_, anal scale;
_b′_, stylet; _c_, support of the stylet; _e_, base or support of
sting (_fi_); 3, profile showing the relation of the genital
armature to the rest of the abdomen, the 6th sternite having been
drawn to show its full size; 4, anal scale (_b_) and stylet; _e_,
_i_, supports and body of the stylet; _c_, piece uniting the two
scales; 5, lateral scale (_a_), and _a_′ sheath; _d_, support of the
sting (_f_); 6, transverse section of the body through the sting
(diagrammatic); _R_, internal armature; _o_, oviduct; _a_, lateral
scale; _a′_ its valve; _e_, support of the stylets (_i_); _b_, anal
scale; _c_, piece uniting two scales; _f_, sting; _d_, its support;
7, a second section simpler and more theoretical than the first; 8,
diagrammatic, all the elements of the sting have been reduced to
pieces of the same form.—After Lacaze-Duthiers.
]

An objection to this view is the fact that the posterior pairs of styles
appear to arise both from one and the same segment,—the ninth. Dewitz
questions whether the four appendages of the ninth segment represent two
pairs of limbs, or one pair split into two branches, and prefers the
latter view, but leaves it as a point to be settled by future
investigations. As will be seen below, both Kraepelin and Bugnion
observed a pair of rudiments to each of the three penultimate segments,
those of the middle pair splitting in two. Wheeler maintains,
erroneously we think, that the inner of the two pairs on the ninth
segment represents the tenth pair of abdominal appendages; but in
reality this latter pair become the cercopods. That there are probably
originally in insects of all the orders provided with an ovipositor
three distinct pairs of appendages, one to each segment, is proved, or
at least strongly suggested, by Ganin’s researches on the three pairs of
abdominal imaginal discs of the third larva of Platygaster and Polynema
(Fig. 188), which are transformed into the ovipositor. He remarks that
these imaginal discs have the same origin and pass through the same
changes as those in front, _i.e._ those destined to form the thoracic
legs. Dewitz has shown that the germs of the ovipositor of the honey-bee
arise as buds on the two segments before the last (Fig. 189).

[Illustration:

FIG. 188.—Third larva of Polynema: _at_, antenna; _fl_, imaginal buds
of the wing; _l_, of the legs; _tg_, buds of the middle pair of
stylets of the ovipositor; _fk_, fat-body; _eg_, ear-like
process.—After Ganin.
]

[Illustration:

FIG. 189.—Imaginal buds and papillæ of the ovipositor of the honey-bee
attached to tracheæ; at different stages: _b′_, 1st; _b″_, 2d or
middle; and _c′_, 3d pair of papillæ.—After Dewitz.
]

Kraepelin also detected in the larva of the honey-bee a pair of what he
regarded as genuine imaginal buds on abdominal segments eight, nine, and
ten; the buds on the tenth segment are divided each in two; of these
four appendages the two median ones form the barbed sting (_gorgeret_ or
_stachelrinne_), and the two lateral stylets, the valves
(_stachelscheiden_). The two buds of the ninth segment give rise to the
vagina and to the oviducts, and these unite secondarily with the
posterior end of the ovaries. The genital appendages of the male
correspond to those of the female, and arise from four imaginal buds
situated on the under side of the tenth abdominal segment.

[Illustration:

FIG. 190.—1, sting and poison sac of the honey-bee: _GD_, poison
gland; _Gb_, poison reservoir; _D_, accessory gland; _sh_, sheathing
style or sting-“feeler”; _Str_ sting; _Ba_, sheath; _Q_, quadrate
plate; _O_, oblong piece; _W_, angular piece; _B_, base of the sting
and stylets; _Stb′_, _Stb″_, the two barbed stylets or darts. 2,
sting seen from the ventral face: lettering as in the other
figure.—After Kraepelin, from Perrier.
]

In the ants, according to Dewitz, the genital armature is derived from
imaginal buds situated on the under side of the seventh, eighth, and
ninth abdominal segments. Bugnion has observed the formation of six
imaginal buds of the genital armature in the larva of a chalcid
(Encyrtus, Figs. 41, 42, 191, _q^1_, _q^2_, _q^3_), the transformation
of the central part of these structures into small digitiform pads, then
the division of the two intermediate buds into four (?) (Fig. 191, _B_,
_q^2_), but was unable to trace their farther development.

The subject still needs farther investigation, since certain
observers, as Haase, and, more recently, Heymons, do not believe
that they are homologues of the legs, but integumental structures,
though of somewhat higher value than the style of the base of the
legs of Scolopendrella and Thysanura; but it is to be observed that
as yet we know but little of the embryological history of these
styles.

Those authors who have examined the elements of the ovipositor, and
regard them as homologues (_homodynamous_) of the limbs, are
Weismann (1866), Ganin (1869), Packard (1871), Ouljanin (1872),
Kraepelin, Kowalevsky (1873), Dewitz (1875), Huxley (1877),
Cholodkowsky, Bugnion (1891), and Wheeler (1892).

As shown, then, by our observations and those of Dewitz (Figs. 189 and
192), the rudiments of the ovipositor consist of three pairs of
tubercles, arising, as Kraepelin and also Bugnion (Fig. 191) have shown,
from three pairs of imaginal discs, situated respectively on the
seventh, eighth, and ninth uromeres, or at least on the three
penultimate segments of the abdomen. With the growth of the semipupa,
the end of the abdomen decreases in size, and is gradually incurved
toward the base (Fig. 193), and the three pairs of appendages approach
each other so closely that the two outer ones completely ensheath the
inner pair, until a complete extensible tube is formed, which, by the
changes in form of the muscles within, is gradually withdrawn entirely
within the body.

[Illustration:

FIG. 191.—_A_, end of larva of Encyrtus of 2d stage, showing the three
pairs of imaginal buds of the ovipositor _q^1_, _q^2_, _q^3_. _B_,
the same in an older larva ready to transform; _i_, intestine; _x_,
genital gland; _a_, anus.—After Bugnion.
]

An excellent account of the honey-bee’s sting is given by Cheshire
(Figs. 194, 195). The outermost of the three pairs of stylets
forming the apparatus is the two thick, hairy “palpi” or feelers
(_P_), these being freer from the sting proper than in the
ovipositor of Orthoptera. The sting itself is composed of the two
inner pairs of stylets; one of these pairs is united to form the
sheath (_sh_), while the other pair form the two barbed darts. The
sheath has three uses: first, to open the wound; second, to act as
an intermediate conduit for the poison; and third, to hold in
accurate position the long barbed darts. The sheath does not enclose
the darts as a scabbard, but is cleft down the side presented in
Fig. 194, which is below when the sting points backward. But, says
Cheshire, as the darts move up and down, they would immediately slip
from their position, unless prevented by a mechanical device,
exhibited by _B_ and _C_, giving in cross-section sheath and darts
near the end, and at the middle of the former. “The darts (_d_) are
each grooved through their entire length, while upon the sheath
(_sh_) are fixed two guide rails, each like a prolonged dovetail,
which, fitted into the groove, permits of no other movement than
that directly up and down.” The darts are terminated by ten barbs of
ugly form (_D_, Fig. 194), and much larger than those of the sheath,
and as soon as the latter has established a hold, first one dart and
then the other is driven forward by successive blows. These in turn
are followed by the sheath, when the darts again more deeply plunge,
until the murderous little tool is buried to the hilt. But these
movements are the result of a muscular apparatus yet to be examined,
and which has been dissected away to bring the rigid pieces into
view. The dovetail guides of the sheath are continued far above its
bulbous portion, as we see by _E_, Fig. 195; and along with these
the darts are also prolonged upward, still held to the guides by the
grooved arrangement before explained; but both guides and darts, in
the upper part of their length, curve from each other somewhat like
the arms of a Y, to the points _c_, _c′_ (_A_, Fig. 194), where the
darts make attachment to two levers (_i_, _i′_). The levers (_k_,
_l_ and _k′_, _l′_) are provided with broad muscles, which terminate
by attachment to the lower segments of the abdomen. These, by
contraction, revolve the levers aforesaid round the points _f_,
_f′_, so that, without relative movement of rod and groove, the
points _c_, _c′_ approach each other. The arms of the Y straighten
and shorten, so that the sheath and darts are driven from their
hiding-place together and the thrust is made by which the sheath
produces its incision and fixture. The sides being symmetrical, we
may, for simplicity’s sake, concentrate our attention on one, say
the left in the figure. A muscular contraction of a broad strap
joining _k_ and _d_ (the dart protractor) now revolves _k_ on _l_,
so that _a_ is raised, by which clearly _c_ is made to approach _d_;
_i.e._ the dart is sent forward, so that the barbs extend beyond the
sheath and deepen the puncture. The other dart, and then the sheath,
follow, in a sequence already explained, and which _G_, Fig. 195, is
intended to make intelligible, _a_ representing the entrance of the
sheath, _b_ the advance of the barbs, and _c_ the sheath in its
second position. The barb retractor muscle is attached to the outer
side of _i_, and by it _a_ is depressed and the barbs lifted. These
movements, following one another with remarkable rapidity, are
entirely reflex, and may be continued long after the sting has been
torn, as is usual, from the insect. By taking a piece of
wash-leather, placing it over the end of the finger, and applying it
to a bee held by the wings, we may get the fullest opportunity of
observing the sting movements, which the microscope will show to be
kept up by continued impulses from the fifth abdominal ganglion and
its multitudinous nerves (_n_, Fig. 194, _A_), which penetrate every
part of the sting mechanism, and may be traced even into the darts.
These facts, together with the explanation at page 49, will show why
an abdomen separated many hours may be able to sting severely, as I
have more than once experienced.

[Illustration:

FIG. 192.—Base of the ovipositor of _Locusta viridissima_ seen from
beneath: _c′_, sheath, or outer and lower pair of stylets turned to
one side to show the others; _b′_, upper and inner pair; _b″_, third
or innermost, smallest pair of stylets. _A_, the same on one side,
in section. The shaded parts show the muscular attachments. The
muscles which extend the apparatus and are attached to ν, δ, and η,
as also the membranes which unite the pieces from η; to γ with each
other and the body, are removed, so that only the chitinous parts
remain.—After Dewitz.
]

[Illustration:

FIG. 193.—Development of the sting in Bombus: _A_, _a_, 1st pair on
8th sternite; _b_, 2d inner pair forming the darts; _c_, outer pair.
_B_-_E_, more advanced stages. _F_, _x_, _y_, _z_, three pairs of
tubercles, the germs of the male organs.
]

[Illustration:

FIG. 194.—Sting of bee × 30 times: _A_, sting separated from its
muscles; _ps_, poison sac; _pg_, poison gland; _5th g_, 5th
abdominal ganglion; _n_, _n_, nerves; _e_, external thin membrane
joining sting to last abdominal segment; _i_, _k_, _l_, and _i′_,
_k′_, _l′_, levers to move the darts; _sh_, sheath; _v_, vulva; _p_,
sting-palpus or feeler, with tactile hairs and nerves. _B_ and _C_,
sections through the darts and sheath, × 300 times: _sh_, sheath;
_d_, darts; _b_, barbs; _p_, poison-channel. _D_, end of a dart, ×
200: _o_, _o_, openings for poison to escape into the wound.—After
Cheshire.
]

[Illustration:

FIG. 195.—Details of sting of bee: _E_, darts, sheath, and valves;
_pb_, poison-bag duct; _fo_, fork; _s_, slide piece; _va_, valve;
_b_, barbs. _F_, terminal abdominal segments; _w_, worker’s sting;
_q_, queen’s sting; _r_, _r′_, anal plate; _G_, sting entering skin;
_sh_, sheath; _a_, _b_, _c_, positions in first, second, and third
thrusts with the sting. _H_, portion of poison gland, × 300; _cn_,
cell nucleus; _n_, nerve; _g_, ganglionic cell. _I_, portion of the
poison gland, cells removed; _cd_, central duct; _d_, individual
small ducts; _pr_, tunica propria. _K_, gland of _Formica rufa_;
_cd_, central duct; _d_, small ducts; _sc_, secreting cells. _L_,
valve and support; _t_, trachea; _va_, valve; _tr_, truss or
valve-prop.—After Cheshire.
]

[Illustration:

FIG. 196.—_A_, rudimentary ovipositor of nymph of Æschna. _B_, the
corresponding ♂ structures; _a_, enlarged. _C_, ovipositor of nymph
of Agrion; _d_, gill.
]

The male genital armature in the bees is originally composed of three
pairs of tubercles, homologous with those of the female, all originally
arising from three abdominal segments, two afterward being anterior, and
the third pair nearer the base of the abdomen.

The ovipositor of the dragon-flies (Odonata) is essentially like that of
the Orthoptera and Hymenoptera. Thus in Æschna (Fig. 196), Agrion (Fig.
196, _C_), and also in Cicada it consists of a pair of closely appressed
ensiform processes which grow out from under the posterior edge of the
eighth uromere and are embraced between two pairs of thin lamelliform
pieces of similar form and structure.

=The styles and genital claspers= (_Rhabdopoda_).—Other appendages of
the end of the abdomen of pterygote insects, and generally, if not
always, arising from the ninth segment, are the clasping organs, or
_rhabdopoda_ as we may call them, of Ephemeridæ (Fig. 197), Neuroptera
(Corydalus [Fig. 198], Myrmeleon, Rhaphidia), Trichoptera, Lepidoptera,
Diptera, and certain phytophagous Hymenoptera. They do not appear to
occur in insects which are provided with an ovipositor. In Thysanura the
styles are present on segments 1–9 (Fig. 179). Those of the male
Ephemeridæ, of which there are two pairs arising from the ninth segment,
are remarkable, since they are jointed, and they serve to represent or
may be the homologues of two of the pairs of stylets composing the
ovipositor of insects of other orders. The lower pair (Fig. 197, _rh_)
are either 2–, 3–, or 4–jointed (in Oniscigaster 5–jointed), while those
of the upper pair are 2–jointed (_rh′_). These rhabdopods in the
ephemerids are evidently very primitive structures, since they approach
nearest in shape and in being jointed to the abdominal legs of
Scolopendrella and the Myriopoda. The styles of the Orthoptera are
survivals of the embryonic appendages of the ninth segment (Wheeler,
etc.). In Mantis they are seen to have the same relations as the cerci,
as shown by Heymons (Fig. 200).

[Illustration:

FIG. 197.—Abdomen of _Ephemera_ (Leptophlebia) _cupida_, ♂: _c_, base
of cercopoda; _rh_, outer 3–jointed claspers or rhabdopods; _rh′_,
inner pair. _A_, side view.
]

[Illustration:

FIG. 198.—End of abdomen of _Corydalus cornutus_, ♂: _vh_, rhabdopod;
_c_, cercopod.
]

In the Phasmidæ, in Anabrus, and in the Odonata the cercopods, which are
not jointed, are converted into claspers, and in the Odonata the
claspers are spiny within, so as to give a firmer hold. The suranal
plate is apparently so modified as to aid in grasping the female. In
nearly all the Trichoptera there are, besides the suranal plate, which
is sometimes forked (Nosopus), a pair of superior and of inferior
claspers, and in certain genera (Ascalaphomerus, Macronema, Rhyacophila,
Hydropsyche, Amphipsyche, Smicridea, and Ganonema) the lower pair are
2–jointed like those of Ephemeridæ. The number of abdominal segments in
the adult Trichoptera is nine, and McLachlan states that the genital
armature consists of three pairs of appendages, _i.e._ a superior,
inferior, and intermediate pair, besides the suranal plate (vestige of a
tenth segment) and the penis. Judging by his figures, these three pairs
of appendages arise from the last or ninth uromere, and the upper pair
seem to be the homologue of the cercopoda of ephemerids. It needs still
to be ascertained whether the intermediate pair is a separate set, or
merely subdivisions of the upper or lower, and whether one of the latter
may not arise from the penultimate segment, because we should not expect
that the last segment should bear more than one pair of appendages, as
we find to be the case in arthropods in general, and in the Neuroptera,
from which the Trichoptera may have originated.

[Illustration:

FIG. 199.—End of abdomen of embryo of Mantis: _r_, rhabdopod; _c_,
cercopod; _sp_, suranal plate; _st_, stigma on 8th segment.—After
Heymons.
]

[Illustration:

FIG. 200.—End of abdomen of _Periplaneta americana_, ♂, side view:
_c_, cercopod; _st_, stilus; _p_, penis; _t_, titillator; _d_,
“bird’s head” (clasper?); _i_, “oblong plate”; IX-XI, terminal
segments; X, suranal plate; XI′, 11th sternite.—After Peytoureau.
]

In most larvæ of the Trichoptera, especially the Rhyacophilidæ and
Hydropsychidæ, the last abdominal segment bears a pair of 2–jointed legs
(cercopoda), ending in either one or two claws, which under various
forms, sometimes forming long processes, persist in the pupa; and there
appears to be a suranal plate, the vestige of the tenth uromere. In the
pupa, judging by Klapálek’s figure of Leptocerus (24_{8,9}, 25), a pair
of lateral spines arise which may in the imago form one of the pairs of
appendages or styles. In the pupa of _Œcetis furva_ his figure 28_{9}
shows two pairs of 1–jointed appendages arising from the last segment;
whether the long dorsal or upper styles arise from the vestige of a more
distal segment is not distinctly shown in Klapálek’s sketch. The origin
of these elements of the genital armature evidently needs further study.

Whether the abdominal legs or so-called false or prop-legs of
lepidopterous larvæ are genuine legs, homologous with those of the
thorax and with the cephalic appendages, or whether they are secondary
adaptive structures, is a matter still under discussion. That, however,
they are true legs is shown by the embryology of the Lepidoptera, where
there is a pair to each abdominal segment. It may also be asked whether
the anal legs of lepidopterous larvæ are not the homologues of the
2–jointed anal appendages of caddis-worms.

[Illustration:

FIG. 201.—_Eriocephala calthella_, ♂, side view: _t_, palpiform
suranal plate; _cl_, claspers; _s_, inferior claspers; _mxp_,
maxillary palpi; _cx_. coxa; _tr_, trochanter; _sc_, scutum; _sc′_,
scutellum.
]

In Lepidoptera, notably the male of the very generalized _Eriocephala
calthella_ (Fig. 201), besides the broad unjointed claspers, which are
curved upward and provided with a brush of stiff hooked setæ (this upper
pair being perhaps modified cercopods), there is an accessory lower
slenderer pair, while the suranal plate (_t_) is palpiform or clavate
and also adapted to aid in the action of the claspers. The examination
of the cercopods and rhabdopods in the Trichoptera and in a generalized
lepidopterous form like this enables one to understand the morphology of
the genital armature, since it consists, besides the suranal plate,
which is often deeply forked (in Sphingidæ, Smith), of a pair of
modified hook-like cercopoda, and in some cases (Eriocephala) of an
additional pair of claspers which may be the homologues of the ephemerid
rhabdopods. A pair of hooks, often strong and claw-like (_harpes_), are
situated, one near the base on the inside of each clasper; they are
especially developed in the Noctuidæ (Smith), and appear to be present
in certain Trichoptera, but this remains to be proved. This complicated
apparatus of claspers and hooks is utilized by those insects which pair
while on the wing, and is wanting in such forms as Coleoptera and
Hemiptera. Besides the forceps of Panorpa, there are two pairs of
slender filiform appendages which need farther examination. In the
Diptera, especially Tipulidæ, there is a pair of 2–jointed appendages or
forceps, as in Limnophila (Osten Sacken). The male genital armature of
Diptera appears to be on the same general plan as in Lepidoptera, but
more complicated.

[Illustration:

FIG. 202.—Male organs of generation of Athalia.—After Newport.
]

Notice should also be taken of the paired uncinate hooks which are
modifications of the penis-sheath of the male of cockroaches
(Phyllodromia), which Haase states appear to originate on the tenth
ventral plate, and which probably “serve to open and dilate the vagina
of the female, especially as a perforated penis, which is highly
developed in Machilis, seems to be wanting in the Blattidæ.” (Haase.)

=The penis.=—This is a single or double median style-like structure
either hollow and perforated, or solid, very variable in shape,
receiving the end of the ejaculatory duct. It is usually enclosed
between two lateral plates, the homologues perhaps of the inner pair of
sheaths of the ovipositor. In the Coleoptera, as in Carabidæ and
Melolonthidæ, the penis is a long chitinous tube, “retractile within the
abdomen on the under surface as far as the anterior segments.”
(Newport.) In the Hymenoptera, of which that of the saw-flies is a type,
Newport states that it “consists of a short valvular projectile organ,
covered externally by two pointed horny plates (_i_) clothed with soft
hairs.” Above these are two other irregular double-jointed plates (Fig.
202, _l_) surrounded at their base by a chitinous ring (_k_); they are
edged with prehensile hooked spines (_i_). Between these in the middle
line are two elongated muscular parts (_m_) which enclose the penis
(_h_), and which, like those in beetles, perhaps aid in dilating the
vulva of the female.

An examination of Figs. 203–207 will aid in understanding the various
modifications in beetles, etc., of this organ.

A general study of the anatomy and homologies of the male genital
armature, from a developmental point of view, together with a comparison
of them with the corresponding female organs, is still needed.

[Illustration:

FIG. 203.—End of abdomen situated under the anal lobes of _Hydrophilus
piceus_, drawn out, seen from the ventral side: 6, sternal region of
6th segment; 7, 8, 9, segments telescoped, when retracted, in 6th
segment; _zw_, membrane connecting 6th and 7th segments; _G_,
intromittent apparatus; _vl_, external lobes; _vlu_, inner lobes;
_pn_, penis.
]

[Illustration:

FIG. 204.—The same as in Fig. 203, seen from the side: 6, the free 6th
segment; 7–10, the four last, when at rest, retracted and telescoped
within the 6th segment, with the copulatory apparatus (_g_); _vl_,
outer, _vlu_, inner lobe; 10_s_, tergite of 10th segment; 10_i_,
sternite of the same; _an_, anal opening.
]

[Illustration:

FIG. 205.—Terminal parts of the male copulatory apparatus of
_Hydrophilus piceus_, torn apart: _vlu_, the two inner lodes; _pn_,
penis; _x_, membrane torn from under side of penis; _ej_,
ejaculatory duct; _os_, its opening on the under side of the penis,
directly under its tip. The muscles, tracheæ, and nerves are not
drawn.
]

=Velum penis.=—In the locusts (Acrydiidæ) the penis is concealed by a
convex plate, flap, or hood, free anteriorly and attached posteriorly
and on the sides to the ridge forming the upper edge of the tenth
sternite. When about to unite sexually, the tip of the abdomen is
depressed, the hood is drawn backward, uncovering the chitinous penis.

=The suranal plate.=—This is a triangular, often thick, solid plate or
area, the remnant of the tergum of the last, usually tenth, segment of
the abdomen, the supra-anal or suranal plate, or anal operculum (_lamina
supra-analis_) of Haase. In most lepidopterous larvæ this plate is well
marked; in those of the Platypteridæ it is remarkably elongated, forming
an approach to a flagellum-like terrifying appendage, and in that of
_Aglia tau_ it forms a long, prominent, sharp spine. In the cockroach,
both Cholodkowsky and Haase maintain that the tenth abdominal segment is
suppressed in the female, the tergal portion being fused with the
suranal plate (the latter in this case, as we understand it, being the
remnant of the eleventh segment of the embryo). As to the nature of the
middle jointed caudal appendage in Thysanura and May-flies Heymons has
satisfactorily shown that it is a hypertrophied portion of the suranal
plate, being in Lepisma but a filamental elongation of the small
eleventh abdominal tergite.

[Illustration:

FIG. 206.—Copulatory organ of a weevil, _Rhychophorus phœnicis_, seen
from above. _A_, _vl_, the lobes united into a capsule; _pp_, torn
membrane which connects the capsule with the 9th abdominal segment;
_ej_, ejaculatory duct. _B_, the same seen from the side; _mu_, end
of the muscle of the penis. _C_, the same as _B_, without the
capsule; _os_, opening of the ejaculatory duct (_ej_). Other letters
as in _A_.
]

[Illustration:

FIG. 207.—_A_, penis (_pn_) of _Carabus hortensis_: _bl_, wrinkled
membranous vesicle; _vlu_, the valves; _g_, part of 9th segment.
_B_, end of penis of the same, enlarged; _os_, cleft-like opening;
also a wrinkled vesicle, as at _bl_.—This and Figs. 203–205 after
Kolbe.
]

At the base of the suranal plate of locusts (Acrydiidæ) is the suranal
fork or suranal furcula (_furcula supra-analis_, as we have called it)
(Fig. 88, 89, _f_).

=The podical plates or paranal lobes.=—In the cockroach and other
insects, also in the nymphs of Odonata, the anus is bounded on each side
by a more or less triangular plate, the two valves being noticeable in
lepidopterous larvæ. They are the _valvulæ_ of Burmeister, and podical
plates of Huxley, who also regarded them as the tergites of an eleventh
abdominal segment;[38] and the subanal laminæ of Heymons. They are
wanting in Ephemeridæ.

=The infra-anal lobe.=—Our attention was first called to this lobe or
flap, while examining some geometrid larvæ. It is a thick, conical,
fleshy lobe, often ending in a hard, chitinous point, and situated
directly beneath the vent. Its use is evidently to aid in tossing the
pellets of excrement away so as to prevent their contact with the body.
The end may be sharp and hard or bear a bristle. Whether this lobe is
the modified ventral plate of the ninth urite, we will not undertake at
present to say.

=The egg-guide.=—In the Acrydiidæ the external opening of the oviduct is
bounded on the ventral side by a movable, triangular, acute flap, the
egg-guide (Fig. 88, _B_, _eg_). Whether this occurs in other orders
needs to be ascertained.

LITERATURE ON THE ABDOMEN AND ITS APPENDAGES

_a._ General (including the cerci, stili, etc.)

=Cornelius, C.= Beiträge zur naheren Kenntnis von _Palingenia
longicauda_. (Programm d. Real- u. Gewerbeschule zu Elberfeld, 1848,
pp. 1–38, 4 Taf.)

=Schiödte, J. G.= Bemerkungen über Myrmecophilen. Ueber den Bau des
hinterleibes bei einigen Käfergattungen. (Germar’s Zeits. f. Ent.,
1844.)

—— De metamorphosi Eleutheratorum observationes. (Naturhist. Tidsskr.
i-xiii, 1861–1883.)

=Meinert, F.= Anatomia Forficularum. Anatomisk undersogelse af de
Danske Orentviste, i. (Naturhist. Tidsskr. 3 raekke, ii, 1863, pp.
427–482, 1 Pl.)

—— Om dobbelte Saedgange hos insecter. (Naturhist. Tidsskrift, 3
raekke, v, 1868.)

=Tullberg, Tycho.= Sveriges Podurider. (Svenska Vetensk.-Akad. Handl.,
1872, x, Nr. 10, 12 Pls.)

=Davis, H.= Notes on the pygidia and cerci of insects. (Journ. R.
Micr. Soc., 1879, ii.)

=Westhoff, F.= Ueber den Bau des Hypopygiums der Gattung Tipula Meig.
(Münster, 1882, pp. 1–62, 6 Taf.)

=Saunders, Edward.= Further notes on the terminal segments of aculeate
Hymenoptera. (Trans. Entom. Soc. London, 1884, pp. 251–267, 1 Pl.)

=Palmén, J. A.= Ueber paarige Ausführungsgange der Geschlechtsorgane
bei Insekten. (Helsingfors, 1884, 5 Taf.)

=Haase, E.= Die Abdominalanhänge der Insekten mit Berücksichtigung der
Myriopoden. (Morpholog. Jahrbuch, 1889, xv, pp. 331–435, 2 Taf.)

—— Abdominalanhänge bei Hexapoden. (Sitzungsber. d. Gesellsch.
naturforsch. Freunde, 1889, pp. 19–29.)

=Wheeler, William M.= On the appendages of the first abdominal segment
of embryo insects. (Trans. Wis. Acad. Sc., viii, 1890, pp. 87–140, 3
Pls.)

=Janet, Charles.= Études sur les fourmis, 5^e note, sur la morphologie
du squelette des segments post-thoraciques chez les Myrmicides.
(Mém. Soc. Acad, de l’Oise, xv, pp. 591–611, Figs. 1–5, 1894.)

=Heymons, Richard.= Die Segmentirung des Insektenkörpers. (Anh. Abh.
Akad. Berlin, Phys. Abh., pp. 39, Taf., 1895. See also Die
Embryonalent, von Dermapteren und Orthopteren, under Embryology.)

—— Grundzüge der Entwicklung und des Körperbaues von Odonaten und
Ephemeriden. (Abhandl. k. Preuss. Akad. d. Wissens. Berlin, 1896, 2
Taf., pp. 1–66.)

—— Zur Morphologie der Abdominalanhänge bei den Insecten. (Morphol.
Jahrb., iv, 1896, pp. 178–203, 2 Taf.)

=Peytoureau, S. A.= Contribution à l’étude de la morphologie de
l’armure génital des insectes. (Bordeaux, 1895, pp. 248, 22 Pls., 43
Figs. in text.)

=Verhoeff, C.= Cerci und styli der Tracheaten. (Ent. Nachr., xxi, pp.
166–168, 1895.)

Also the writings of Oudemans, Packard, etc.

_b._ The ovipositor

=Lacaze-Duthiers, Henri.= Recherches sur l’armure génitale femelle des
insectes. (Ann. d. sc. natur., 1849, xii, pp. 353–374, 1 Pl.; 1850,
xiv, pp. 17–52, 1 Pl. (Hyménoptères); 1852, xvii, pp. 206–251, 1 Pl.
(Orthoptères); 1853, xix, pp. 25–88, 4 Pl. (Neuroptères, Thysanures,
Coléoptères, Diptères); pp. 203–237 (Lépidoptères, Aphaniptères en
general. Also separate.))

=Sollmann, A.= Der Bienenstachel. (Zeitschr. f. Wissens. Zool., xiii,
1863, pp. 528–540, 1 Taf.)

=Fengger, H.= Anatomie und Physiologie des Giftapparates bei den
Hymenopteren. (Archiv f. Naturgesch., 1863, pp. 139–178, 1 Taf.)

=Eaton, A. E.= Remarks upon the homologies of the ovipositor. (Trans.
Ent. Soc. London, 1868, pp. 141–144.)

=Packard, A. S.= On the structure of the ovipositor and homologous
parts in the male insects. (Proc. Boston Soc. Nat. Hist., xi, 1868,
pp. 393–399, Figs. 1–11.)

=Lambrecht, A.= Samtliche Teile des Stechapparates in Bienenkörper und
ihre Verwendung zu technischen und vitalen Zwecken. (Bienemwirtsch.
Centralbl., 7 Jahrg., 1871, pp. 5–11.)

=Kraepelin, K.= Untersuchungen über den Bau, Mechanismus und die
Entwicklung des Stachels der bienenartigen Tiere. (Zeitschr. f.
wiss. Zoologie, xxiii, 1873.)

=Dewitz, H.= Vergleichende Untersuchungen über Bau und Entwicklung des
Stachels der Honigbiene und der Legescheide der grünen Heuschrecke.
(Königsberg, 1874.)

—— Ueber Bau und Entwicklung des Stachels und der Legescheide einiger
Hymenopteren und der grünen Heuschrecke. (Zeitschr. f. wissen.
Zoologie, xxv, 1874, pp. 174–200, 2 Taf.)

—— Ueber Bau und Entwicklung des Stachels der Ameisen. (Zeitschr. f.
wiss. Zoologie, 1877, xxviii, pp. 527–556, 1 Taf.)

=Adler, H.= Legeapparat und Eierlegen der Gallwespen. (Deutsche Entom.
Zeitschr., 1877, xxi Jahrg., pp. 305–332, 1 Taf.)

=Cholodkowsky, N.= Ueber den Hummelstachel und seine Bedeutung für die
Systematik. (Zool. Anzeiger, vii, 1884, pp. 312–316.)

=Ihering, H. von.= Der Stachel der Meliponen. (Ent. Nachr., 1886, xii,
Jahrg., pp. 177–188, 1 Taf.)

=Meinert, F.= Bidrag til de danske Myrers Naturhistorie. (Kjöbenhavn,
1890, 68 s.u. Danske Vidensk. Selsk. Skrifter, 5 Raek, v, 3 Pls.)

=Beyer, O. W.= Der Giftapparat von _Formica rufa_ ein reduziertes
Organ. (Jena. Zeitschr. Naturw., 1890, xxv, pp. 26–112, 2 Taf.)

=Carlet, G.= Mémoire sur le venin et l’aiguillon de l’abeille. (Ann.
d. sc. nat. Zool., 7 sér., ix, 1890, pp. 1–17, 1 Pl.)

=Künckel d’Herculais, J.= Méchanisme physiologique de la ponte chez
les insectes orthoptères de la famille des Acridides.—Rôle de l’air
comme agent mécanique et fonctions multiples de l’armure génitale.
(Compt. Rend., cxix, pp. 244–247, 1894.)

Also the writings of Verhoeff, Heymons.

_c._ The external genital armature

=Klug, Johann C. F.= Versuch einer Berichtigung der Fabriciusschen
Gattungen Scolia u. Tiphia. (Ueber u. Mohr Beiträge zur Naturkunde,
i, pp. 8–40, 1805, 1 Taf.)

=Audouin, J. V.=, and =Lachat=. Observations sur les organes
copulateurs males des Bourdons. (Annal. général, d. sc. phys., 1821,
viii, pp. 285–289.)

=Audouin, J. V.= Lettre sur la génération des insectes. (Ann. des sc.
nat., sér. 1, ii, 1824.)

=Suckow, F. W.= L. Geschlechtsorgane der Insekten. (Heusinger,
Zeitschr. organ. Physik., 1828, ii, pp. 231–264, 1 Taf.)

=Rathke, H.= De Libellularum partibus genitalibus. Regiomonti, 1832,
pp. 6 + 38, 3 Pls.

=Siebold, C. Th. E. von.= Ueber die Fortpflanzungsweise der
Libellulinen. (Germar’s Zeitschr. f. Ent., 1840, ii, pp. 421–438.)

=Selys-Longchamps, E. de.= Monographie des Libellulidees d’Europe. 4
Pls., 1840.

=Bassi, C. A.= Studi sulle funzioni degli organi genitali degl’
insetti da lui osservati piu specialmente nella _Bombyx mori_. (Atti
della 5 Riun. d. Scienz. Ital. Lucca, 1844, pp. 39–94.)

=Stein, F.= Vergleichende Anatomie und Physiologie der Insekten, i.
Die weiblich. Geschlechtsorgane der Käfer. Berlin, 1847, 9 Taf.

=Ormancey, P.= Recherches sur l’étui penial considéré comme limite de
l’espèce dans les coléoptères. (Ann. sc. nat., 1849, 3 sér., Zool.,
xii, pp. 227–242.)

=Roussel, C.= Recherches sur les organes génitaux des insectes
coléoptères de la famille des Scarabéides. (Compt. rend. Acad. d.
sc. Paris, 1860, l, pp. 158–161.)

=MacLachlan, R.= A monographic revision and synopsis of the
Trichoptera of the European fauna. London, 1874–80, 59 Pls.

—— On the sexual apparatus of the male Acentropus. (Trans. Ent. Soc.
London, 1872, pp. 157–162.)

=Thomson, C. G.= Nagra anmarkningar ofver arterna af slagtet Carabus.
(Thomson’s Opuscula Entomologica, vii, 1857, pp. 615–729, 1 Pl.)

=Dufour, L.= Sur l’appareil génital male du _Coræbus bifasciatus_.
(Thomson, Archiv ent., 1857, i, pp. 378–381.)

=White, F. Buchanan.= On the male genital armature in the Rhopalocera.
(Trans. Linn. Soc., ser. 1, Zool., i, pp. 357–369, 1876, 3 Pls.)

=Graber, V.= Die Aehnlichkeit im Baue der ausseren weiblichen
Geschlechtsorgane bei den Lokustiden und Akridiern auf Grund ihrer
Entwicklungsgeschichte. (Sitzber. k. Akad. d. Wissensch. Wien.,
1870, lxi, pp. 1–20, 1 Taf.)

=Scudder, Samuel H., and Edward Burgess.= On asymmetry in the
appendages of hexapod insects, especially as illustrated in the
lepidopterous genus Nisoniades. (Proc. Boston Soc. Nat. Hist., 1871,
xiii, pp. 282–306.)

=Chadima, J.= Ueber die Homologie zwischen den männlichen und
weiblichen ausseren Sexualorganen der Orthoptera Saltatoria Latr.
(Mitteil. d. naturwiss. Vereins f. Steiermark, 1872, pp. 25–33, 1
Taf.)

=Hagens, von.= Ueber die Genitalien der männlichen Bienen, besonders
der Gattung Sphecodes. (Berlin Ent. Zeitschr., 1874, pp. 25–43.)

—— Ueber die männlichen Genitalien der Bienengattung Sphecodes.
(Deutschen Entom. Zeitschr., 1882, pp. 209–228, 2 Taf.)

=Lindenman, C.= Vergleichend-anatomische Untersuchung ueber das
männliche Begattungsglied der Borkenkäfer. (Bull. Soc. Imp. d.
Natural. Moscou, 1875–77.)

=Forel, A.= Der Giftapparat und die Analdrüsen der Ameisen. (Zeitschr.
f. wiss. Zoologie, xxx, suppl., 1878.)

=Kraatz, G.= Ueber die Wichtigkeit der Untersuchung des
männlichen Begattungsgliedes der Käfer für die Systematik und
Artunterscheidung. (Deutschen Entom. Zeitschr., 1881, xxv, pp.
113–126.)

—— Ueber das männliche Begattungsglied der europaischen Cetoniiden und
seine Verwendbarkeit für deren scharfe spezifische Unterscheidung.
(Ibid., pp. 129–149.)

=Gosse, Ph. H.= On the clasping-organs ancillary to generation in
certain groups of the Lepidoptera. (Trans. Linn. Soc., 1882, Ser. 2,
Zool., ii, pp. 265–345, 8 Pls.)

—— The prehensors of male butterflies of the genera Ornithoptera and
Papilio. (Proc. Roy. Soc. London, 1881, xxxiii, pp. 23–27.)

=Radoszkowski, O.= Revision des armures copulatrices des males du
genre Bombus. (Bull. Soc. Natur. Moscou, 1884, xlix, pp. 51–92, 4
Pls.)

—— Revision des armures copulatrices des males de la tribu
Philérémides. (Ibid., 1885, lxi, pp. 359–370, 2 Pls.)

—— Revision des armures copulatrices des males de la famille des
Mutillidæ. (Horæ Soc. Ent. Ross., 1885, xix, pp. 3–49, 9 Pls.)

—— Revision des armures copulatrices des males de la tribu des
Chrysides. (Horæ Soc. Ent. Ross, xxiii, 1890, pp. 3–40, 6 Pls.)

=Hofmann, O.= Beiträge zur Kenntnis der Butaliden. (Stett. Ent. Zeit.,
1888, pp. 335–347, 1 Taf.)

=Driedzichi, H.= Revue des espèces européennes du genre Phronia Winn.
(Horæ Soc. Ent. Ross., 1889, xxiii, pp. 404–532, 10 Pls.)

=Sharp, David.= On the structure of the terminal segment in some male
Hemiptera. (Trans. Ent. Soc. London, 1890, pp. 399–427, 3 Pls.)

=Escherich, K.= Die biologische bedeutung der Genitalanhänge der
Insekten. (Verhandl. d. zool. bot. Ges. Wien., 1892.)

—— Anatomische Studien über das männliche Genitalsystem der
Coleopteren. (Zeits. f. wissens. Zool., lvii, pp. 620–641, 1 Taf., 3
figs.)

=Verhoeff, C.= Zur vergleichenden Morphologie der “Abdominalanhänge”
der Coleopteren. (Ent. Nachr., xx, pp. 93–96, 1894. Compare also O.
Schwarz and J. Weise’s criticisms in D. Ent. Zeit., pp. 153–157;
also pp. 101–109, 155–157. Zool. Anzeiger, pp. 100–106, 1894.)

—— Vergleichende—morphologische Untersuchungen ueber das Abdomen
der Endomychiden, etc., und über die Musculature des
Copulationsapparates von Triplax. (Archiv f. Naturg., lxi, pp.
213–287, 2 Taf., 1895.)

—— Vergleichende Untersuchungen über die abdominal Segmente der
weiblichen Hemiptera-Heteroptera und Homoptera. (Verh. Nat. Ver.
Bonn, l, pp. 307–374, 1894.)

=Verhoeff, C.= Beiträge zur vergleichenden Morphologie des Abdomens
der Coccinelliden, etc. (Archiv f. Naturg., lxi, pp. 1–80, 6 figs.,
1895.)

=Boas, J. E. V.= Organe copulateur et accouplement der hanneton.
(Oversigt over det K. Danske Vidensk. Selskab Forhand, 1892,
Copenhagen, 1893, 1 Pl., pp. 239–261.)

=Pérez, J.= De l’organe copulateur mâle des Hyménoptères et de sa
valeur taxonomique. (Ann. Soc. Ent. France, lxiii, pp. 74–81, Figs.,
1894.)

=Goddard, Martha Freeman.= On the second abdominal segment in a few
Libellulidæ. (Proc. Amer. Philos. Soc., xxxv, pp. 205–212, January
11, 1897, 2 Pls.)

Also the writings of Eaton, Emery, Fischer, Forel, Géhin, Godart,
Hagen, Joly, Koletani, Loew, Meinert, Mik, Nicolet, Osten Sacken,
Pictet, Roussel, Schaeffer (1754), Schaum, Schenk, J. B. Smith,
Thompson, Buchanan-White, Brunner von Wattle-Wyll, Weise, Wyenbergh.

The subject of copulation has been treated by Hoffer, Hartig,
Schiedeknecht, Verhoeff, etc.

THE ARMATURE OF INSECTS: SETÆ, HAIRS, SCALES, TUBERCLES, ETC.

[Illustration:

FIG. 208.—Larva of _Dryocampa rubicunda_, stage II.—Bridgham _del._
]

=The cuticula.=—The integument is externally either smooth and shining
or variously punctured, granulated, tuberculated, striated, or hairy. In
certain orders the skin is clothed with flattened setæ or scales, while
many forms, as some caterpillars (Figs. 208, 209), beetles (Fig. 210),
etc., are protected by spines, horns, etc., these in adult insects often
forming secondary sexual characters, usually being more developed in the
males than in the females.

[Illustration:

FIG. 209.—Larva of _Hyperchiria io_, on hatching.
]

The cuticula is not always smooth, but is often finely granulated or
even minutely spinulated. On the abdominal segments of Anabrus, as
observed by Minot, the cuticula is armed with microscopic conical
nodules scattered irregularly over it. They do not correspond, he
says, in any way to hairs; for they do not rest over pores, nor did
he see any specially modified cells underlying them. “As far as I
have observed, they are mere local irregularities, each nodule being
apparently supported by some four or six unmodified epidermal
cells.” Minot adds that the whole of the cuticula, except the cones
just described and the hairs, is divided into numerous minute
fields, each of which corresponds to a single cell of the underlying
hypodermis. Each field is bounded by a distinct polygonal outline,
and its surface is either covered by a large number of extremely
minute projecting points, as on the dorsal arch of the segment, or
is smooth, as upon the articular membrane and ventral arch. Upon the
sides of the dorsal arch and upon the spiracular membrane each field
has a projecting spine or sometimes two or even three. (See also pp.
28, 30.)

[Illustration:

FIG. 210.—_Phanæus pegasus_, ♂, from Mexico.—After Graber.
]

[Illustration:

FIG. 211.—Section of integument of _Datana ministra_: _c_, cuticula;
_hyp_, hypodermis; _p_, outer pigmented nodulated layer.
]

The cuticle of lepidopterous larvæ has also been described and
figured by Minot. In the caterpillars of different groups
investigated by him, the cuticle was found to be rough with
microscopic teeth or spinules, erect or flattened and scale-like,
and either densely crowded or scattered, and affording excellent
generic and specific characters. In the slug-worms (Limacodids) we
have observed that the cuticula is unusually rough, especially on
the spiniferous tubercle of Empretia, Parasa, etc. (Fig. 213, _c_).
The skin of the body between the tubercles is seen to be finely
shagreened, due to the presence of fine teeth, which are more or
less curved and bent, these teeth arising from a very finely
granulated surface (_d_). The cuticle of neuropterous,
trichopterous, and tenthredinid larvæ will probably afford similar
cases. The integument of the larva of Datana is, on the black bands,
rough and nodulated, the irregular nodules being filled with a black
pigment, and forming a layer (_p_) external to the true cuticula
(Fig. 211).

[Illustration:

FIG. 212.—Hairs of Datana: _f_, formative hair-cell; _c_, cuticula;
_p_, pigmented layer; _hy_, hypodermis.
]

The integument of many insects contains fine canals passing through the
chitinous layers and opening externally in minute pores. Certain of the
pore-canals communicate with hollow setæ which sit directly over the
pores; other pores form the external openings of dermal glands, but in
many cases they are empty or only filled with air, and do not have any
hairs connected with them. Each of these pores communicates with a
hair-forming hypodermal cell, called by Graber a _trichogen_.

=Setæ= (“hairs” and bristles).—The setæ of insects are, as in worms,
processes of the cuticle originating from certain of the hypodermal
cells. They arise either from a ring-like pit, or from a minute
tubercle, and are usually situated at the outlet of a pore-canal, which
connects with an underlying cell of the hypodermis (Fig. 212). They are,
then, bristle or hair-like processes arising from the hypodermis. Where
the hairs or setæ are rubbed off, their site is indicated by a minute
ring like a follicle in the chitinous integument. The cuticular hair,
says Leydig, is in its first condition the secretion of the cellular
element of the skin, and a thread-like continuation of the cell-body may
rise up through the pore-canal into the centre of the hair, remaining
there permanently.

While the setæ are usually simple, they are often branched, plumose, or
spinulose, as in larval Hemerobiidæ, Anthrenus, and Dermestes, the larvæ
of certain coccinellid beetles, notably Epilachna, and of Cassida, the
larvæ of arctians, etc., and in bees (Anthophila, Megachile, Osmia,
Colletes, Apis, etc.).

The use of these spinulose, plumose, and twisted hairs in the bees is
clearly shown by J. B. Smith, who states that as these insects walk over
flowers, the pollen grains adhere to the vestiture, “and this also
accounts for the fact, probably noticed by every observant fruit-grower,
that bees frequently bury themselves completely in the blossoms, or roll
over every part of them. Such insects are after pollen, not honey, and
by so rolling about, the pollen grains are brought into contact with and
adhere to the surface of the insect.” The syrphid flies also pollenize
flowers, the pollenizing of chrysanthemums being effected, as Smith
states, by _Eristalis tenax_, and he adds that the body vestiture of the
syrphids “is often composed of spurred and branched hairs.” (For
reference to gathering hairs, see p. 45.)

[Illustration:

FIG. 213.—Cuticular spinules of larva of Adoneta: _a_, _b_, _c_, _d_,
different forms; _e_, _e′_, caltrops.
]

Certain remarkable spines occur in limacodid larvæ, notably Empretia and
Adoneta. These we have called caltrops spines, from their resemblance to
the caltrops formerly used in repelling the attacks of cavalry. They are
largely concerned in producing the poisonous and irritating effects
resulting from contact with the caterpillars of these moths, and are
situated in scattered groups near the end of the tubercles. A group of
three is represented at Fig. 213, _e_. They are not firmly embedded in
the cuticle, but on the contrary appear to become very easily loosened
and detached, and they probably, when brought into contact with the skin
of any aggressor, burrow underneath, and are probably in part the cause
of the continual itching and annoyance occasioned by these creatures. It
will be seen by reference to Fig. 213, _e′_, that the body of the spine
is spherical, with one large, elongated, conical spine arising from it,
the spherical base being beset with a number of minute, somewhat obtuse
spinules.

[Illustration:

FIG. 214.—Glandular hairs of caterpillars. _A_, _Dasylophia anguina_:
_a_, of body; _b_, of head; _c_, of prothoracic shield. _B_,
_Ceratosia tricolor_: _a_, on body; _b_, on abdominal legs. _C_,
_Schizura ipomeæ_: _a_, from third thoracic segment; _b_, from larva
stage II; _c_, simple setæ from minute warts.
]

=Glandular hairs and spines.=—In some insects occur fine, minute, hollow
setæ from which exude, perhaps through pore-canals of extreme fineness,
droplets of a clear watery or plasma-like sticky fluid. The club-shaped
tenent hairs of the feet of Collembola, and the hairs fringing the feet
of Diptera, are modified glandular hairs. Here they serve to give out a
sticky fluid enabling the insect to walk on smooth surfaces; they end in
a vesicle-like bulbous expansion, which may contain numerous
pore-canals. Those of caterpillars were first noticed by Zeller, and
Dimmock has particularly described those of the larvæ of Pterophoridæ.
They are either club-shaped, or variously forked at the end (Fig. 214,
_B_, _a_). They are usually replaced after the first larval moult by
ordinary, simple, solid, pointed setæ, and their use in caterpillars is
as yet unknown. Whether these hairs, as seems most probable, arise from
a specialized glandular hypodermal cell, or not, has not yet been
discovered.

[Illustration:

FIG. 215.—_A_, group of setæ arising from a subdorsal tubercle: _cut_,
the cuticle; _hy_, the hypodermis; _sc_, the enlarged and
specialized cells of the hypodermis which secrete the spines
themselves; _pglc_, the nuclei which secrete the venomous fluid
which fills the cavity of the seta (_s_), seen at _p_ in a broken
spine. _B_, a short entire, and a long broken seta (_s-p_); _pgle_,
four poison cells; _p_, the poison in the hollow of the spine.
]

These temporary fine glandular hairs are probably the homologues of the
larger true glandular bristles and spines of the later stages of certain
lepidopterous larvæ, which are brightly colored and lead an exposed
life, living through a large part of the summer. In these structures the
bristles or spines are hollow, filled with a poisonous secretion formed
in a single large, or several smaller specialized hypodermal cells
situated under the base of the spine. In the venomous spines of _Lagoa
crispata_ the poisonous fluid in the larger spines (Figs. 215, _C_, 216,
_b_) is secreted in several large cells situated at the base of the
spine, and this is the usual form. In the finer spines of a large
tubercle (Figs. 215, _A_, 216) there appears to be a differentiation of
the hypodermal cells into two kinds, the large, basal deep-seated,
setigenous cells (216, _sc_) and the poison-secreting nuclei (216,
_pglc_) situated nearer the base of the setæ. The spines being filled
with poison and breaking into bits in the skin of the hands or neck,
cause great irritation and smarting. These nettling or poisonous hairs
or spines are especially venomous in the larva of Orgyia, _Empretia
stimulea_, _Hyperchiria io_, the larvæ of the saturnians (Fig. 217) and
lasiocampids, etc. They rarely occur in insects of other orders, though
the skin of Telephorus is said by Leydig to bear glandular hairs.

[Illustration:

FIG. 216.—Section of a subdorsal tubercle from a larva in stage 1:
_sc_, the setigenous cells, one for each seta; _pglc_, nuclei by
which the poison is secreted; _s_, seta; _p_, poison in middle of a
broken spine; _cut_, cuticle; _sd_, _tub_, spinulated surface of the
subdorsal tubercle.
]

Leydig states that in the stout bristles of Saturnia there is, as in
the integument of the body, a homogeneous cuticula, under which is
the cellular matrix (hypodermis), and the clear contents
(hyaloplasma) are secreted from the blood. The cell-structure of the
hairs consist, as in the cells of the body, of spongioplasma and
hyaloplasma. Leydig has observed the droplets of the secretion of
the caterpillar of _Saturnia carpini_ oozing through distinctly
observable pores, and states that there are similar openings in the
hairs and scales. Dewitz found easily observable openings at the end
of the hair of a large exotic weevil (Fig. 130).

The advanced nymph of Psylla is also armed with clavate glandular
hairs (Fig. 178).

[Illustration:

FIG. 217.—Armature of last four segments of _Callosamia promethea_:
_a_, a dorsal seta; _b_, one showing the poison (_p_) within.
]

The tubercles are outgrowths of the body-walls; they are either
smooth, warty, or spiny, as in many caterpillars. While the armature
of insects is of little morphological importance, it is evidently of
great biological importance, the welfare or even the life of the
insect depending upon it; and it varies in each species of insect,
especially in Diptera, where the position of even a single seta
characterizes the species.

[Illustration:

FIG. 218.—Section through an antennal pectination of _Saturnia
carpini_: _a_, hypodermis, formative cells of the hairs (_c_); _d_,
cuticula; _e_, trachea.—After Semper.
]

[Illustration:

FIG. 219.—Flattened hairs from the lateral tufts of larva of
_Gastropacha americana_: _A_, three from the lateral tuft of
_Heteropacha rileyana_.
]

The mode of development of the hairs was first described by Semper. In
the pectination of the antenna of _Saturnia carpini_ he observed that
the hairs arise, like the scales of the wings, from large round
formative-cells lying in the cavity, which send out through the
hypodermis and cuticle a long slender process which finally becomes the
hair (Fig. 218).

Tactile hairs are those setæ arising over nerve cells or nerve
terminations and will be discussed under the organs of sense.

[Illustration:

FIG. 220.—The same in _G. quercifolia_: _a_, a small hair ending in
two minute processes.
]

=Scales.=—In very rare cases the hairs of caterpillars (Fig. 219) are
flattened and scale-like, and this passage in the same insect of
cylindrical hairs into flattened scale-like ones, shows that the scales
are only modified hairs. Also, as we shall see farther on, Semper has
proved that their mode of origin is identical. While true scales are
characteristic of Synaptera (Thysanura and Colembola), as well as
Lepidoptera and Trichoptera, they also occur in the Psocidæ
(Amphientomum), in many Coleoptera (Curculionidæ, Cleridæ, Ptinidæ,
Dermestidæ, Byrrhidæ, Scarabæidæ, Elateridæ, and Cerambycidæ), and in
the Culicidæ, and a few other Diptera, though they are especially
characteristic of the Lepidoptera, not a species of this great order
being known to be entirely destitute of them.

[Illustration:

FIG. 221.—Flattened and spinulated hairs of tufts of larva of
_Acronycta hastulifera_.
]

[Illustration:

FIG. 222.—Scales from dorsal tuft, on second thoracic segment of larva
of _Gastropacha quercifolia_.
]

The scales vary much in shape, but are more or less tile-like, attached
to the surface of the body or wing by a short slender pedicel, and are
more loosely connected with the integument than the hairs, which are
thicker at the base or insertion than beyond.

The markings of the scales, both of Synaptera and Lepidoptera, are very
elaborate, consisting of raised lines, ridges, or striæ with transverse
ridges between. “The striæ of the transparent scales of Micropteryx are
from about 500 to 300 to the millimetre, varying in different species.
The opaque scales of Morpho, which show metallic reflections, have about
1400 striæ to the millimetre.” (Kellogg.)

The primary use of scales, as observed by Kellogg, is to protect the
body, as seen in Synaptera and Lepidoptera. A nearly as important use is
the production of colors and patterns of colors and markings, while in
certain butterflies certain scales function as the external openings of
dermal scent-glands, and they afford in some cases (as first claimed by
Kettelhoit in 1860) generic and specific characters. Spuler has shown
that the scales are strengthened by internal chitinous pillars. Burgess
has observed in the scales of _Danais plexippus_ that the under surface
of the scales is usually smooth, or provided with few and poorly
developed ridges, and this has been confirmed by Spuler and by Mayer
(Fig. 226).

In the irised and metallic scales the ridges, says Spuler, are not
divided into teeth, and they converge at the base to the pedicel and
also toward the end of the scale (Micropteryx), or end in a single
process beyond the middle (the brass-colored scales of _Plusia
chrysitis_).

The arrangement of the scales on the wings is, in the generalized moths,
irregular; in the more specialized forms they are arranged in bands
forming groups, and in the most specialized Lepidoptera they are more
thickly crowded, overlapping each other and inserted in regular rows
crossing the wings, these rows either uniting with each other or running
parallel. (Spuler.) The scattered irregular arrangement seen in
Micropteryx is also characteristic of the Trichoptera and of
Amphientomum.

[Illustration:

FIG. 223.—Portion of a longitudinal section through one of the young
pupal wings of a summer pupa of _Vanessa antiopa_: _s_, young scale;
_leu. cy._, leucocyte; _mbr. pr._, ground membrane; _prc_,
hypodermis-cells.
]

[Illustration:

FIG. 224.—Portion of a longitudinal section through one wall only of
the pupal wing of a specimen slightly older than that of Fig. 223;
_s_, older scale.
]

=Development of the scales.=—The mode of origin of the scales was first
worked out by Semper in 1886, who stated that in the wing of the pupal
Sphinx and Saturnia they are seen, in sections, to arise from large
roundish cells just under the hypodermis and which have a projection
which passes out between the hypodermis (his “epidermis”) cells,
expanding into a more or less spherical vesicle, the latter being the
first indication of the future scale. He observed that the scales are
not all formed at once, but arise one after another, so that on one and
the same wing the scales are in different stages of development.

[Illustration:

FIG. 225.—Portion of a longitudinal section through a pupal wing about
eight days before emergence: _s_, formative scale-cell; upper _s_, a
scale.
]

More recently Schaeffer has stated that the scales and also the hairs
are evaginations of greatly enlarged hypodermis cells, and still more
complete evidence has been afforded by A. G. Mayer (1896). In the wings
of Lepidoptera, about three weeks before the imago emerges, certain of
the hypodermis cells, which occur at regular intervals, begin to
increase in size and to project slightly above the level of the
hypodermis; these are Semper’s “formative cells,” and are destined to
secrete the scales. They increase in length, and appear as in Fig. 223.
In the next stage observed, the projections are much longer (Fig. 224).
The hypodermis is now thrown up into a regular series of ridges, which
run across the wing. Each ridge, says Mayer, corresponds in position
with a row of formative cells, and each furrow with the interval between
two adjacent rows. The scales always project from the tops of these
ridges. The ground or basal membrane has not participated in this
folding, and the deep processes of the hypodermis (_prc_) that once
extended to this membrane have largely disappeared. Figure 225
represents a more advanced stage almost eight days before the emergence
of the imago.

The scales are originally filled with protoplasm, which gradually
withdraws, leaving behind it little chitinous bars or pillars which
serve to bind together the upper and lower surfaces of the scales, and
finally the scales become “merely little flattened hollow sacs
containing only air.” As Mayer shows (Figs. 226, 227), from the study of
scales examined four days before emergence of the butterfly (Danais),
“the striations upon the upper surface of the scale are due to a series
of parallel longitudinal ridges,” while the under side is usually
smooth.

The mode of insertion is seen in Fig. 227. The narrow cylindrical
pedicel of the scale is merely, according to Semper, inserted into a
minute close-fitting socket, which perforates the wing-membrane, and not
into a tube, as Landois supposed. Spuler describes a sort of double sac
structure or follicle (_Schuppenbalg)_ which receives the hollow pedicel
of the scale. This was originally (1860) observed by F. J. Carl Mayer,
but more fully examined by Spuler (Fig. 228) though not detected by A.
G. Mayer.

[Illustration:

FIG. 226.—Portion of a cross-section through the pupal wing of _Danais
plexippus_, about six days before emergence: _sg_, scale; _cta.al_,
wing-membrane; _cl.frm_, formative cell of the scale; _mbr.pr_,
ground-membrane; _fbr.h′drm_, hypodermal fibres of pupal wings. _A_,
portion of a longitudinal section through the pupal wing, eight or
nine days before emergence; _prc_, processes of young hypodermis
scales.—This and Figs. 223–225 after Mayer.
]

=Spinules, hair-scales, hair-fields, and androconia.=—Besides the
scales, fine spinules occur on the thickened veins of the wings of the
Blattidæ, where they resemble fir-cones; also in the Perlidæ, in the
Trichoptera, and in the more generalized Lepidoptera (Micropterygidæ and
Hepialidæ), occur, as indicated by Spuler, delicate chitinous hollow
spinules scarcely one-tenth as long as, and more numerous than, the
scales, which sometimes form what he calls “Haftfelds,” or holding
areas. These spinules have also been noticed by Kellogg, and by myself
in Micropteryx; Kellogg, and also Spuler, have observed them in certain
Trichoptera (Hydropsyche). These also occur on the veins, and detached
ones near large one-jointed hairs, or hair-scales, said by Kellogg to be
striated. Kellogg has detected these scale-hairs, as he calls them, in
Panorpa.

[Illustration:

FIG. 227.—View looking down upon the upper (_i.e._ exposed) surface of
one of the large scales situated on the veins of _Danais plexippus_,
about four days before emergence: _clm_, chitinous pillars found in
scales. _A_, _a_ smaller scale, _a_, _a′_, sections of the scales.
_B_, leucocyte found in the larger scale.—After Mayer.
]

[Illustration:

FIG. 228.—Scale-follicles: _A_, of a scale of _Galleria mellonella_:
_r_, neck-ring. _B_, the same of _Polyommatus phlæas_. _C_, the same
of a hair on inner edge of hind wing of _Lycæna alexis_ ♀.—After
Spuler.
]

[Illustration:

FIG. 229.—_A_, portion of wing of a caddis-fly (Mystacides). _B_,
enlarged, showing the androconia and hair-scales. _C_, a separate
androconium.—After Kellogg.
]

The “hair-scales” of the phylogenetically older Trichoptera correspond
to certain scales of Lepidoptera, especially the Psychidæ (Spuler),
variously called “plumules” (Deschamps), “battledore scales,” also
certain minute cylindrical hairs. To these scent-scales is applied the
term _androconia_. They are found, almost without exception, on the
upper side of the fore wings, occurring in limited areas, such as the
discal spots, or on folds of the wings. Fritz Müller has shown that they
function as scent-scales, and are confined to the males. Kellogg has
detected androconia-like scales on the wings of a caddis-fly,
_Mystacides punctata_ (Fig. 229).

[Illustration:

FIG. 280.—Cross-section of androconia surface on wing of _Thecla
calanus_; _a_, androconia; _gl_, gland of base; _s_, ordinary
scales; _w_, wing in section.—After Thomas.
]

Thomas has proved by sections of the wing of Danais, etc., that the
androconia arise from glands situated in a fold of the wing (Fig. 230),
and he states that the material elaborated by the local glands, and
distributed upon the surface of the wing by the androconia, is that
which gives to many of the Lepidoptera their characteristic odor. On
comparing these “glands,” it is evident that they are groups of
specialized formative cells of Semper (trichogens), which secrete an
odorous fluid, issuing perhaps from extremely fine pore-canals at the
ends of the androconia. They thus correspond to the glandular hairs,
poison-hairs, and spines of caterpillars, the formative cells of which
contain either a clear lymph or poison.

LITERATURE

_a._ Hairs, bristles, cleaning spines, calcaria, combs, etc.

=Leydig, Franz.= Zum feineren Bau der Arthropoden. (Müller’s Archiv f.
Anat. und Phys., 1855, pp. 376–480.)

=Fobel, Auguste.= Les fourmis de la Suisse. Bâle, 1874.

=Saunders, Edward.= Remarks on the hairs of some of our British
Hymenoptera. (Trans. Ent. Soc. London, 1878, pp. 169–171.)

=Perez, J.= Notes d’apiculture. (Bull. Soc. d’Apic. de la Gironde,
Bordeaux, 1882.)

=Osten Sacken, C. R. von.= An essay on comparative chætotaxy, or the
arrangement of characteristic bristles of Diptera. (Trans. Ent. Soc.
London, 1884, pp. 497–517.)

—— Preliminary notice of a subdivision of the suborder Orthorrhapha
Brachycera (Diptera) on chætotactic principles. (Berlin Ent.
Zeitschr., 1896, pp. 365–373.)

=Janet, Charles.= Études sur les fourmis. 8^e Note. Sur l’organe de
nettoyage tibio-tarsien de _Myrmica rubra_. (Ann. Soc. Ent. France,
1895, pp. 691–704, 6 Figs.)

See also J. B. Smith’s Economic Entomology, 1896, hairs of bees. Also
the writings of De Geer, Huber, Fenger, Mayr, Forel, Canestrini, and
Berlese (1880); Dahl, Cheshire, etc.

_b._ Glandular and poisonous setæ and spines

=Ratzeburg, J. Th. Ch.= Ueber entomologische Krankheiten. (Stettin
Ent. Zeit., 1846, vii, pp. 35–41.)

=Zeller, P. C.= Revision der Pterophoriden. (Linnæa Ent., vi, pp.
319–416, 1852, at p. 356 speaks of “Drüsenhärchen.”)

=Dimmock, George.= On some glands which open externally on insects.
(Psyche, iii, pp. 387–401, 1882.)

=Goossens, Th.= Des chenilles urticants. (Ann. Soc. Ent. France, 1881,
pp. 231–236.) Des chenilles vésicants. (Ibid., 1886, pp. 461–464.)

=Packard, A. S.= Notes on some points in the external structure and
phylogeny of insects. (Proc. Boston Soc. Nat. Hist., xxv, 1890, pp.
83–114, 2 Pls.)

—— A study of the transformations and anatomy of _Lagoa crispata_, a
bombycine moth. (Proc. Amer. Phil. Soc., xxxii, pp. 275–292, 7 Pls.,
1894.)

=Holmgren, Emil.= Studier öfner hudens och de körtelartade hudorganens
morfologi hos Skandinaviska macrolepidopterlarver. (K. Svenska
Vetenskaps-Akad. Handl., xxvii, pp. 1–83, Stockholm, 1895, 9 Pls.)

Also the writings of Leydig, Keller, Bach, Karsten, Scribner, Riley,
etc.

(See also Literature of repugnatorial glands.)

_c._ Androconia

=Deschamps, Bernard.= Récherches microscopiques sur l’organisation des
ailes der Lépidoptères. (Ann. des Sc. nat. [?], iii, pp. 111–157,
1835.)

=Waufor, T. W.= On certain butterfly scales characteristic of sex.
(London, 1867–68.)

=McIntire, S. J.= Notes on the minute structure of the scales of
certain insects. (London, 1871.)

=Anthony, J.= The markings on the battledore scales of some of the
Lepidoptera. (London, 1872.)

=Scudder, S. H.= Antigeny or sexual dimorphism in butterflies. (Proc.
Amer. Acad. Arts and Sc., xii, 1877, pp. 150–158.) Also Butterflies,
etc. (New York, 1881, pp. 192–206, figs.)

=Müller, Fritz.= A prega costal das Hesperideas. (Archivas do Museo
nac. do Rio de Janeiro, iii, pp. 41–50, 2 Pls., 1878.)

=Thomas, M. B.= The androconia of Lepidoptera. (Amer. Nat., xxvii, pp.
1018–1021, 2 Pls., 1893.)

_d._ Scales

=Leydig, Franz.= Zum feineren Bau der Arthropoden. (Archiv f. Anat.
und Phys., 1855, pp. 376–480, 1 Taf.)

=Semper, Carl.= Beobachtungen über die Bildung der Flügel, Schuppen,
und Haare bei den Lepidopteren. (Zeitschrift f. wissensch. Zoologie,
1857, pp. 326–339, 1 Taf.)

=Mayer, F. T. Karl.= Ueber den Staub der Schmetterlingsflügel.
(Allgem. mediz. Centralzeitung, 1860, pp. 772–774.)

=Landois, H.= Beiträge zur Entwicklungsgeschichte der
Schmetterlingsflügel in der Raupe und Puppe. (Zeitschr. f.
wissensch. Zoologie, xxi, 1871, pp. 305–316, 1 Taf.)

=Weismann, August.= Ueber Duftschuppen. (Zool. Anzeiger, i, 1878, pp.
98–99.)

=Dimmock, George.= Scales of Coleoptera. (Psyche, iv, pp. 1–11, 23–27,
43–47, 63–71, 1883.)

=Schaeffer, Cäsar.= Beiträge zur Histologie der Insekten. (Zool.
Jahrbücher, Abth. f. Anat. u. Ontog., iii, pp. 611–652, 2 Pls.,
1889.)

=Kellogg, Vernon L.= The taxonomic value of the scales of the
Lepidoptera. (Kansas Univ. Quart., iii, pp. 45–89, figs. 1–17, 9
Taf., 1894.)

=Mayer, Alfred G.= The development of the wing-scales and their
pigment in butterflies and moths. (Bull. Mus. Comp. Zool., xxix.,
1896, pp. 209–236, 7 Pls.)

=Spuler, Arnold.= Beiträge zur Kenntniss des feineren Baues und der
Phylogenie der Flügeltedeckung der Schmetterlings. (Zool. Jahrb.
Abth. f. Anat. u. Ontog., viii, pp. 520–543, 1 Taf., 1895.)

—— Ueber das Vorhandensein von Schuppenbalg bei den Schmetterlingen.
(Biol. Centralblatt, xvi, Sept. 15, 1896, pp. 677–679, 3 figs.)

THE COLORS OF INSECTS

The colors and bright markings of insects, especially those of
butterflies, render them the most brilliant and beautiful creatures in
existence, rivalling and even excelling the gay hues of our most
splendidly colored birds. The subject has been but recently taken up and
is in a somewhat crude condition, but the leading features have been
roughly sketched out by the work of a few observers from a physical,
chemical, and biological point of view.

The colors of insects, as of all other animals, are primarily due to the
action of light and air; other factors are, as Hagen observes, heat and
cold, moisture and dryness, as recently shown by the experiments on
butterflies by Dorfmeister, Weismann, W. H. Edwards, and later
observers. They have their seat in the integument. Hagen divides colors
into optical and natural.

=Optical colors.=—“These,” says Hagen, “are produced by the interference
of light, and are by no means rare among insects, but they are solely
optical phenomena. Colors by the interference of light are produced in
two different ways: either by thin superposed lamellæ, or by many very
fine lines or small impressions in very close juxtaposition.

“1. There must be present at least two superposed lamellæ to produce
colors by interference. The naked wings of Diptera, of dragon-flies, and
of certain Neuroptera often show beautiful interference colors. The
wings of Chrysopa and Agrion show interference colors only for a certain
time, viz., as long as the membranes of the wings are soft and not
firmly glued together. Afterwards such wings become simply hyaline.

“The scales of Entimus and other Curculionidæ are well known for
their brilliancy, and it is interesting to remark that when dry
scales are examined with the microscope, many are found partly
injured, which give in different places different colors, according
to the number of layers which remain. The elytra of some
Chrysomelina and other beetles with iridescent colors probably
belong to the same category.

“2. When there are scales with many fine lines or small impressions
close to each other, we have the second mode of producing colors.

“The fine longitudinal and transversal lines of lepidopterous scales
seem to serve admirably well to produce the brilliant effect of
color-changing butterflies. But there must be something more present, as
most of the scales of Lepidoptera are provided with similarly fine
lines, and only comparatively few species change colors. I remark
purposely that the lines in the color-changing scales are not in nearer
juxtaposition.” (Hagen.)

“The colors of butterflies change mostly from purple to blue,
sometimes to yellow. The splendid violet color at the end of the
wings of _Callosune ione_ is brought out by a combination of the
natural with interference colors. Originally the scales are colored
lake-red; but a blue interference color is mixed with it; hence the
violet hue results. The blue tones, _i.e._ the splendid varying blue
of the Morpho butterflies, Schatz claims, owe their hue less to the
interference of light than to a clouded layer of scales situated
over the dark ground, through which the light becomes reflected on
the same. The scales of the Morphids are in reality brown, as we see
by transmitted light; moreover, only the upper side of the scales
sends off blue reflections—the under side is simply brown. But the
blue scales of Urvilliana are also shining blue beneath; by
transmitted light they appear as if clear yellow. The smaragd-green
scales of Priamus show by transmitted light a bright red-orange, and
the orange-yellow of Crœsus a deep grass-green.” (Schatz in Kolbe.)

“Krukenberg presumes the golden-green color of _Carabus auratus_ to
be an interference color. It is not changed by the interference of
light, nor was he able to extract from the elytra any green pigment
with ether, benzol, carbon of sulphur, chloroform, or alcohol, even
after having previously submitted the elytra to the influence of
muriatic acid or ammonia. Chlorophyll is not present, whether free
or combined with an acid.” (Hagen.)

Leydig has shown that the interference colors of the hairs of
certain worms (Aphrodite and Eunice) may be produced by very small
impressions in juxtaposition, which bring about the same effect as
striæ. Such an arrangement occurs on the feathers of birds, _i.e._
on the necks of pigeons and elsewhere, and Hagen suggests that this
kind of interference colors occurs more frequently among insects
than is commonly known. At least the limbs of certain forms appear
yellow, but when held in a certain position change to brown or
blackish. “I know of no other explanation of this not uncommon fact
on the legs of Diptera, of Hymenoptera, and of Phryganidæ.”
Interference colors, he adds, may occur in the same place together
with natural colors. “The mirror spots of _Saturnia pernyi_ show
besides the interference colors a white substance in the cells of
the matrix, which Leydig believes to be guanin. But this fact is
denied by Krukenberg for the same species and also for _Attacus
mylitta_ and _Plusia chrysitis_.”

=Natural colors.=—These are divided by Hagen into _dermal_ (cuticular)
and _hypodermal_. The dermal colors are due to pigment deposited in the
form of very small nuclei in the cuticula. Hagen considers them as
“produced mostly by oxidation or carbonization, in consequence of a
chemical process originating and accompanying the development and the
transformations of insects.”

“To a certain extent the dermal colors may have been derived from
hypodermal colors, as the cuticula is secreted by the hypodermis,
and the colors may have been changed by oxidation and air-tight
seclusion. The cuticula is in certain cases entirely colorless,—so
in the green caterpillar of _Sphinx ocellata_; but the intensely red
and black spots of the caterpillar of _Papilio machaon_ belong to
the cuticula, and only the main yellow color of the body to the
hypodermis.” (Leydig, Histiol., p. 114.)

“The dermal colors are red, brown, black, and all intermediate shades,
and all metallic colors, blue, green, bronze, copper, silver, and gold.
The dermal colors are easily to be recognized as such, because they are
persistent, never becoming obliterated or changed after death.” (Hagen.)

Minot and Burgess refer to the cuticular colors of the cotton-worm
(Aletia), the dark brown color belonging to the cuticula or crust.
“Upon the outside of the crust is a very thin but distinct layer,
which in certain parts rises up into a great number of minute,
pointed spines that look like so many dots in a surface view. Each
spine is pigmented diffusely, and together they produce the brown
markings. The spines are clustered in little groups, one group over
each underlying hypodermal cell.” (U. S. Ent. Comm., 4th Report, p.
46.) Minot also shows that in caterpillars generally a part of the
coloration is caused by pigmentation of the cuticula.

In a dull-colored insect, such as the Mormon cricket (Anabrus), the
coloration, as Minot states, depends principally upon the pigment of
the hypodermis shining through the cuticula. “Most of the cells
contain dull, reddish-brown granules, but scattered in among them
are patches of cells bright green in color. I have observed no cells
intermediate in color; on the contrary, the passage is abrupt, a
brown or red cell lying next a green one. Indeed, I have never seen
any microscopic object more bizarre than a piece of the epidermis of
Anabrus spread out and viewed from the surface.” (2d Report U. S.
Ent. Comm., p. 189.)

The pigment may extend through the entire cuticula, but it is usually
confined to the outermost layers, and occurs there in union with a
peculiar modelling of the upper surface into microscopic figures which
are of interest not only from their delicacy, but because they vary with
each species. (See p. 184.)

The hypodermal colors, situated in the hypodermis, are, according to
Hagen, the result of a chemical process, generating color out of
substances contained in the body. They are easily recognized, since they
fade, change, and disappear after death. But where these colors are
preserved after death and enclosed in air-tight sacs, as in the elytra
and scales and hairs of the body, they persist, though, as we well know,
they may fade after exposure to light.

The hypodermal colors are mostly brighter and lighter than the dermal
ones, being light blue or green in different shades, yellow to orange,
and the numerous shades of these colors combined with white;
exceptionally they are metallic, as in Cassida, and are then obliterated
after death.

“The fact that such metallic colors can be retained in dead
specimens by putting a drop of glycerine under the elytra, leads us
to conclude that those colors are based upon fat substances. The
hypodermal colors are never glossy, as far as I know; the dermal
colors frequently.

“As the wings, elytra, and hairs all possess a cuticula, dermal
colors are frequently to be found, together with hypodermal ones,
chiefly in metallic colors. In the same place both colors may be
present, or one of them alone. So we find hypodermal colors in the
elytra of Lampyridæ. In the elytra of the Cicindelidæ the main
metallic color is dermal, the white lines or spots are hypodermal,
by which arrangement the variability in size and shape of those
spots is explained.

“There occur in a number of insects external colors, that is, colors
upon the cuticula, which I consider to be in fact displaced
hypodermal colors: the mealy pale blue or white upon the abdomen of
some Odonata, the white on many Hemiptera, the pale gray on the
elytra and on the thorax of the Goliath beetle, and the yellowish
powder on Lixus. Some of these colors dissolve easily by ether or
melt in heat, and some of them are a kind of wax. I believe that
those colors are produced in the hypodermis, and are exuded through
the pore-canals.” (Hagen.)

The white colors are simply for the most part due to the inclusion of
air in scales. The white mother-of-pearl spots of Argynnis are produced
by a system of fine transverse pore-canals filled with air; in
Hydrometra the white ventral marks have the same origin. (Leydig.)

The further statements and criticisms of Hagen regarding the relation of
color to mimicry, sexual selection, and the origin of patterns are of
much weight and will be referred to under those heads. Indeed, these
subjects cannot well be discussed without reference to the fundamental
facts stated in the masterly papers of Leydig and of Hagen, and much of
the theorizing of these latter days is ill-founded, because the colors
of insects and animals are attributed to natural selection, when they
seem really the result of the action of the primary factors of organic
evolution, such as changes of light, heat, cold, and chemical processes
dependent on the former.

As to the chemical nature of color, Hagen, after quoting the results of
Krukenberg and others, thinks that the colors of insects are chemically
produced by a combination of fats or fat-acids with other acids or
alkalis under the influence of air, light, and heat. He concludes:—

1. That some colors of insects can be changed or obliterated by acids.

2. That two natural colors, madder-lake and indigo, can be produced
artificially by the influence of acid on fat-bodies.

3. As protein bodies in insects are changed into fat-bodies, and may be
changed by acids contained in insects into fat-acids, the formation of
colors in the same manner seems probable.

4. That colors can be changed by different temperatures.

5. That the pattern is originated probably by a combination of oxygen
with the integument.

6. That mimicry of the hypodermal colors may be effected by a kind of
photographic process.

7. Finally, color and pattern are produced by physiological processes in
the interior of the bodies of insects.

Krukenberg concludes that change of color (in perfectly developed
insects) is a consequence of the change of food, and can be
explained by the alteration of the pigment through heat and light.
His experiments were made in order to ascertain the cause of the
turning of green grasshoppers in autumn into yellow and pink. He
tried to answer two questions: First, does the pigment of
grasshoppers originate directly out of the food, and does it consist
of pure chlorophyll or of a substance containing chlorophyll, or is
it to be accepted as a peculiar product of the organism? Second, is
the color the consequence of only one pigment, or of several?
Special analysis proves that the green color has no connection with
chlorophyll. He concludes: “It is evident that the green color of
the grasshopper is the consequence of several different pigments
which can be separated by a chemical process.” Krukenberg believes
that light has a marked influence on the color of insects and that
light turns to red or pink the insects which were green during the
summer. It would seem, however, more probable that cold was the
agent, the change being due to the colder autumn weather.

Here we might refer to the results of the studies of Buckton and
Sorby, on the changes in color of Aphides:—

“1. The purple coloring matter appears to be a quasi-living
principle, and not a product of a subsequent chemical oxidizing
process. Mounted in balsam or other preserving fluids, the darker
species stain the fluid a fine violet.

“2. As autumn approaches and cold weather reduces the activity of
the Aphides, the lively greens and yellows commonly become converted
into ferruginous red, and even dark brown, which last hue in reality
partakes more or less of intense violet or purple. These changes
have some analogy with the brilliant hues assumed by maple and other
leaves during the process of slow decay.

“3. Aqueous solutions of crushed dark brown and yellow-green
varieties of Aphides originate different colors with acids and
alkalies.

“4. In the generality of cases coloring-matters, such as indigo,
Indian yellow, madder-lake, and the like, do not separately exist in
the substance of vegetables, but the pigments are disengaged through
fermentation or oxygenation. Again, alizarin itself is reddish
yellow, but alkaline solutions strike it a rich violet just as we
find them to act towards the substance which Mr. Sorby calls
aphidilutein.

“5. Mr. Sorby’s four stages of the changes effected by the oxidation
of aphideine produce four different substances.”

=Chemical and physical nature of the pigment.=—Researches in this
difficult field of inquiry have been made by Landois (1864), Sorby
(1871), Meldola (1871), by Krukenberg (1884), and more recently by
Coste, Urech, Hopkins, and Mayer, and the subject is of fundamental
importance in dealing with mimicry and protective coloration, the
primary causes of which appear to be due to the action of physical and
chemical agents.

Over twenty years ago Meldola observed that the yellow pigment of the
sulphur-yellow butterfly (_Gonopteryx rhamni_) was soluble in water, and
showed that its aqueous solution had an acid reaction.

Besides the yellow uranidin found by Krukenberg in different beetles
and lepidopterous pupæ, still other coloring-matters, which are very
constant in different species are readily recognized by the
spectroscope. “Thus there appear in the brownish yellow lymph of
_Attacus pernyi_, _Callosamia promethea_ and _Telea polyphemus_,
after saponification of the precipitated soap readily effected by
ether, or incompletely or not removed by benzine, a chlorophane-like
lipochrome; and in the yellowish green lymph of _Saturnia pyri_ and
of _Platysamia cecropia_ besides this pigment still another whose
spectrum shows a broad band on D, but which disappears with the
addition of acetic acid or ammonia, as also after a long heating of
the lymph up to 66° C.”

Coste, and more especially Urech, have shown that many of the pigments
may be dissolved out of the scales by means of chemical reagents, giving
colored solutions, and leaving the scales white or colorless. They have
also shown that some of these pigments may be changed in color by the
action of reagents, and then restored to their original color by other
reagents. They have proved that reds, yellows, browns, and blacks are
always due to pigments, and in a few cases greens, blues, violets,
purples, and whites, and not, as is usually the case, to structural
conditions, such as striæ on the scales (Mayer). They confined
themselves solely to the chemical side of the problem, not considering
the structure of the scales themselves.

Urech has also discovered a beautiful smaragd-green coloring-matter in
the wings (not in the scales) of the pupa of _Pieris brassicæ_. It is
not chlorophyll, and Urech suggests that it may be either the germinal
substance of the pigments of the scales or its bearer. It is not the
pigment of the blood.

Urech has also demonstrated that in many Lepidoptera the color of the
urine which is voided upon emergence from the chrysalis is similar to
the principal color of the scales.

Hopkins has worked on the pigments within the scales of butterflies.
The yellow pigment in _Gonopteryx rhamni_ is a derivation of uric
acid, and he calls it lepidotic acid. Its aqueous solution is
strongly acid to litmus, and must be bad-tasting to birds.

Hopkins has dissolved the red pigment from the border of the hind
wing of _Delias eucharis_, an Indian butterfly, in pure water,
finding as the result a yellow solution; but if the solution be
evaporated to dryness, the solid residue of pigment is red once
more. He has obtained from this pigment of _eucharis_ a silver
compound which contains a percentage of metals exactly equal to that
from the pigment of _G. rhamni_. (Nature, April 2, 1892.)

“The scales of the wings of the white butterflies (Pieridæ) are also
shown by Hopkins to contain uric acid, this substance practically
acting as a white pigment in these insects. A yellow pigment, widely
distributed in the same family, is shown to be a derivative of uric
acid, and its artificial production as a by-product of the
hydrolysis of uric acid is demonstrated. That this yellow pigment is
an ordinary excretory product of the butterfly is indicated by the
fact that an identical substance is voided from the rectum on
emergence from the pupa. These excretory pigments, which have
well-marked reactions, are apparently confined to the Pieridæ, and
are not found in other Rhopalocera. This fact shows that when a
Pierid mimics an insect belonging to another group, the pigments of
the mimicked and mimicking insects, respectively, are chemically
quite distinct. Other pigments existing, not in the scales, but
between the wing-membranes, are shown to be of use for ornament.”
(Proc. Royal Soc., London, 1894.)

Griffiths (1892) claims that the green pigment found in several
species of Papilio, Hesperia, and Limenitis, also in Noctuidæ,
Geometridæ, and Sphingidæ likewise consists of a derivative of uric
acid, which he calls lepidopteric acid. By prolonged boiling in HCl
it is converted into uric acid.

Spuler, however, finds that green does not depend on pigmentation,
but is an optical color. As remarked by Spuler, either the chitin of
the scales itself is colored reddish (yellow grayish), or the
pigment is secreted in the nuclei.

A. G. Mayer believes that the pigments of the scales are derived from
the hæmolymph or blood of the pupa, for the following reasons: (1) He is
unable to find anything but blood within the scales during the time when
the pigment is formed. (2) In Lepidoptera generally the first color to
appear upon the pupal wings is a dull ochre-yellow, or drab, and this is
also the color assumed by the blood when it is removed from the pupa and
exposed to the air. (3) He has succeeded by artificial means in
manufacturing several pigments from the blood which are similar in color
to various markings upon the wing of the imago; chemical reagents have
the same effect upon these manufactured pigments that they do upon the
similarly colored pigments of the wings. “It should be here noted,” he
says, “that in 1866 Landois pointed out the fact that the color of the
dried blood of many caterpillars is similar to the ground color of the
wings of the mature insect.”

=Ontogenetic and phylogenetic development of colors.=—The colors of the
wings of Lepidoptera, as is well known, are acquired at the end of the
pupal state. The order of development of the colors in the pupal wings
has been observed by Schaeffer, Van Bemmelen, Urech, Haase, Dixey,
Spuler, and A. G. Mayer. The immature wings are at first transparent and
full of protoplasm. The transparent condition of the wings corresponds
to the period before the scales are formed, and when they are full of
protoplasm; they then become whitish as the scales develop; the latter
are at first filled with protoplasm, and afterwards turn whitish, being
little hollow sacks filled with air. After the protoplasm has completely
withdrawn from the scales, the blood of the pupa enters them, and then
the coloring-matter forms. (Mayer.) He adds that “about twenty-four
hours after the appearance of the dull yellow suffusion the mature
colors begin to show themselves. They arise, faint at first, in places
near the centre of the wings, and are distinguished by the fact that
they first appear upon areas between the nervures, never upon the
nervures themselves. Indeed, the last place to acquire the mature
coloration are the outer and costal edges of the wings, and the
nervures.”

The faint color of the scales gradually increases in intensity. “For
example, if a scale be destined to become black, it first becomes pale
grayish brown, and this color gradually deepens into black.”

Urech states that in _Vanessa io_ first a white, and in _V. urticæ_ a
pale reddish hue, are spread over the entire wings, and then
successively arise other colors in the following order: yellow, yellow
to brown, red, brown and black.

Spuler, however, claims that the differentiation of colors and markings
do not follow one another, but arise simultaneously, and that his view
is confirmed by Fischer. This may be the case with the highly
specialized and diversely marked butterflies, but certainly taking the
Lepidoptera as a whole the yellows and drabs must have been the
primitive hues, the other colors being gradually added in the later more
specialized forms.

It is noticeable that the most generalized moths, such as the species of
Micropteryx, Tinea, Psychidæ, Hepialidæ (in general), etc., are dull
brown or yellow-drab without bars, stripes, or spots of bright hues.
These shades prevail in others of the more primitive Lepidoptera, such
as many bombycine moths, and they even appear to a slight extent in
certain caddis-flies. The authors mentioned, especially Mayer, whom we
quote, claim that “dull ochre-yellows and drabs are, phylogenetically
speaking, the oldest pigmental colors in the Lepidoptera; for these are
the colors that are assumed by the hæmolymph upon mere exposure to the
air. The more brilliant pigmental colors, such as bright yellow, reds,
greens, etc., are derived by more complex chemical processes. We find
that dull ochre-yellow and drabs are at the present day the prevalent
colors among the less differentiated nocturnal moths. The diurnal forms
of Lepidoptera have almost a monopoly of the brilliant colorations, but
even in these diurnal forms one finds that dull yellow or drab colors
are still quite common upon those parts of their wings that are hidden
from view.”

The more primitive moths being more or less uniformly yellowish or drab,
the next step was the formation of bars, stripes, finally spots, and
eyed spots, these markings in the later forms appearing simultaneously
in one and the same species of certain highly specialized moths and
butterflies. All that has been said will prepare the reader for the
consideration of the subject of insect coloration. The origin of such
markings has been discussed by Weismann, Eimer, Haase, Dixey, Fischer,
and others.

LITERATURE

=Heer, O.= Einfluss des Alpenklimas auf die Farbe der Insecten.
(Froebel u. Heer, Mitth. aus dem Gebiete der theoret. Erdkunde,
1836, i, pp. 161–170.)

=Goureau.= Mémoire sur l’irisation des ailes des insectes. (Ann. Soc.
Ent. France, 2 sér., i, 1848, pp. 201–215.)

=Laboulbène, A., et M. Follin.= Note sur la matière pulvérulente qui
recouvre la surface du corps des Lixus et de quelques autres
insectes. (Ann. Soc. Ent. de France, 1848, vi, pp. 301–305, Fig.)

=Coquerel, Ch.= Note sur la prétendue poussière cryptogamique qui
recouvre le corps de certains insectes. (Ann. Soc. Ent. France,
1850, viii, pp. 13–15.)

=Brauer, F.= Beobachtungen in Bezug auf den Farbenwechsel bei
_Chrysopa vulgaris_. (Verhandl. k. k. zool.-botan. Gesellsch. Wien.,
1852, pp. 12–14.)

=Prittwitz, O. F. W. v.= Bemerkungen über die geographische
Farbenverteilung unter den Lepidopteren. (Stett. Ent. Zeit., 1855,
xvi, pp. 175–185.)

=Latham, A. G.= The causes of the metallic lustre of the scales on the
wings of certain moths. (Proc. Lit. and Phil. Soc. Manchester, iii,
1864, pp. 198–199. Quart. Journ. Micr. Sc., new ser., iv, 1864, pp.
48–49.)

=Sorby, H. C.= On the coloring matter of some Aphides. (Quart. Journ.
Micr. Sc., new ser., xi, 1871, pp. 352–361.)

=Leydig, Franz.= Bemerkungen über Farben der Hautdecke und Nerven der
Drüsen bei Insekten. (Archiv f. mikr. Anatomie, xii, 1876, pp.
536–550, 1 Taf.)

=Weismann, A.= Studien zur Descendenz-Theorie, ii, 1876.

=Hemmerling, Hermann.= Ueber die Hautfarbe der Insecten. (Bonn, 1878,
p. 27.)

=Buckton, C. B.= Monograph of the British Aphides. (London, 1879, ii,
p. 167.)

=Cameron, P.= Notes on the coloration and development of insects.
(Trans. Ent. Soc. London, 1880, pp. 69–79.)

=Hagen, Hermann A.= On the color and pattern of insects. (Proc. Amer.
Acad. Arts and Sc., 1882, pp. 234–267.)

=Poulton, Edward Bagnall.= The essential nature of the colouring of
phytophagous larvæ (and their pupæ), etc. (Proc. Roy. Soc. London,
xxxviii, pp. 269–315, 1884–1885.)

—— An inquiry into the cause and extent of a special colour-relation
between certain exposed lepidopterous pupæ and the surfaces which
immediately surround them. (Phil. Trans. Roy. Soc. London, clxxviii,
pp. 311–441, 1 Pl., 1887.)

=Krukenberg, C. Fr. W.= Grundzüge einer vergleichenden Physiologie der
Farbstoffe und der Farben. (Heidelberg, 1884, pp. 102.)

=McMunn, C. A.= Krukenberg’s chromatological speculation. (Nature,
xxxi, p. 217, 1885.)

=Müller, Fritz, and Dr. H. A. Hagen.= The color and pattern of
insects. (Kosmos, xiii, 1886, pp. 466–469.)

=Slater, J. W.= On the presence of tannin in insects and its influence
on their colors. (Trans. Ent. Soc. London, 1887, iii, Proceed., pp.
32–34.)

=Bemmelen, J. F. van.= Ueber die Entwicklung der Farben und Adern auf
den Schmetterlingsflügeln. (Tijdschrift der nederland. Dierkundige
Vereeniging, ser. 2, pp. 235–247, 1889.)

=Hopkins, F. G.= Uric acid derivatives functioning as pigments in
butterflies. (Proc. Chem. Soc. London, 1889, p. 117; also Nature,
xl, p. 335.)

—— Pigment in yellow butterflies. (Nature, xlv, p. 197, 1891.)

—— The pigments of the Pieridæ. (Proc. Roy. Soc. London, lvii, No.
340, pp. 5, 6, 1894. Phil. Trans. Roy. Soc. London, clxxxvi, pp.
661–682, 1896.)

=Coste, F. H. P.= Contributions to the chemistry of insect colors.
(The Entomologist, xxiii, 1890; xxiv, 1891, pp. 9–15, etc. Nature,
xlv., pp. 513–517, 541–542, 605.)

=Urech, F.= Beobachtungen über die verschiedenen Schuppenfarben und
die zeitliche Succession ihres Auftretens. (Zool. Anzeiger, xiv, pp.
466–473, 1891; Ibid., August 1, 1892.)

—— Beiträge zur Kenntniss der Farbe von Insektenschuppen. (Zeits. f.
Wissens. Zool., lvii, pp. 306–384, 1893.)

=Griffiths, A. B.= Recherches sur les couleurs de quelques insectes.
(C. R. Acad. Sc. Paris, cxv, pp. 958, 959.)

=Mayer, Alfred Goldsborough.= On the color and color-patterns of moths
and butterflies. (Proc. Bost. Soc. Nat. Hist., xxvii., March, 1897,
pp. 243–330, 10 Pls. See also p. 201 under Mayer.)

Also the writings of Bates, Beddard, Belt, Butler, Darwin, Dimmock,
Dixey, Eimer, Haase, Higgins, Müller, Poulton, Seitz, Wallace,
Weismann.

2. INTERNAL ANATOMY

THE MUSCULAR SYSTEM

In its general arrangement the muscular system of insects corresponds to
the segmented structure of the body. Of the muscles belonging to a
single segment, some extend from the front edge of one segment to that
of the next behind it, and others to the hinder edge; there are also
sets of dorsal and ventral muscles passing in an oblique or vertical
course (Figs. 16–18). As Lang observes, “the greater part of the muscles
of the body can be traced back to a paired system of dorsal and ventral
intersegmental longitudinal muscles.” The muscular system is simplest in
larval insects, such as caterpillars, where the musculature is serially
repeated in each segment.

In the larva of Cossus Lyonet found on one side of the body 217 dorsal,
154 lateral, 369 ventral, and in the thoracic legs 63, or 803 muscles in
all. “Adding to this number the 12 small muscles of the second segment,
and 8 others of the third, which he did not describe, there would be for
all the muscles on one side of the caterpillar 823. This would make for
the entire body 1646, without counting a small single muscle which
occurs in the subdivision of the last segment,” and also those of the
internal organs as well as those of the head, so that the total number
probably amounts to about 2000, not 3000, as usually stated in the
books. Lubbock admits that Lyonet was right in his mode of estimating
the number. In the larva of _Pygærci bucephala_ he found that “the large
muscles scarcely vary at all,” though certain smaller ones are very
variable. Lubbock observed that certain of the longitudinal muscles in
the caterpillar of Diloba split up into numerous, not less than ten,
separate fascicles. “This separation of the fibres composing a muscle
into separate fascicles is carried on to a much greater extent in the
larvæ of Coleoptera. Of course in the imago the number of thoracic
muscles is greatly increased, or at least in Dyticus and the
wood-feeding Lamellicorns, which alone I have examined. In these two
groups each of the larger muscles is represented by at least twenty
separate fascicles, which makes it far more difficult to distinguish the
arrangement of the muscles.”

The muscles are whitish or colorless and transparent, those in the
thorax being yellowish or pale brown; and of a soft, almost gelatinous
consistence. In form they are simply flat and thin, straight, band-like,
or in rare cases pyramidal, barrel or feather shaped. They act variously
as rotators, elevators, depressors, retractors, protractors, flexors,
and extensors.

[Illustration:

FIG. 231.—Diagram of the muscles and nerves of the ventral surface of
the segments in the larva of _Sphinx ligustri_: _A_, _A_, recti
muscles; 1, 2, ventral recti muscles (1, recti majores; 2, recti
minores); 3, ridge giving origin to recti muscles of one segment,
and insertion to the same of the adjoining segment; 4, ridge for
attachment of muscle; 5, retractor ventriculi, connecting the
mid-intestine with the outer integument of the body. _B_, 6, first
oblique,—7, second oblique,—9, 10, third oblique, muscles; 11,
fourth oblique,—12. third rectus,—13, fifth oblique,—14,
triangularis, muscle; 15, transversus medius; 16, transverse ridge;
17, transversi abdominales; 18, abdominales anteriores; 19, 20,
abdominales laterales, some (20) longer than others; 21, obliquus
posterior; 22, postero-laterales obliqui; 23, transversus lateralis;
24, second transversus lateralis; 25, retractor spiraculi, or
constrictor of the spiracles, attached by a long tendon (26); 27,
retractor valvulæ.

Nerves: _a_, ganglion,—_c_, transverse nerves, of which _p_ is the
first, _q_ the second, _r_ the third,
and _s_ the fourth branch; _t_, the main trunk, which crosses the
great longitudinal trachea, receives a
filament from the transverse nerve (_n_), and divides into two
branches (_t_);—some of these branches
form a small plexus (_u_); the nerve _t_ divides in two divisions (_p_
and _v_). The second division ends
in _w_ and _x_; the branch _q_ divides into _y_ and _z_. For other
explanations, see Newport, art. _Insecta_.—After
Newport.
]

[Illustration:

FIG. 232.—Musculature of the European cockchafer, _Melolontha
vulgaris_: _a_, _a_, levatores capitis; _b_, depressores capitis;
_c_, rotatores capitis; _d_, depressors externi; _e_, retractor or
flexor of the jugular plate; _f_, oblique extensor of the jugular
plate; _g_, the other retractor of the jugular plate; _h_, retractor
prothoracis superior; _i_, inferior retractor, the proper depressor
of the prothorax; _k_, elevator prothoracis; _l_, one of the
rotatores prothoracis; _m_, _n_, _o_, flexors of the coxa; _x_,
great depressor muscle of the wing; _y_, _y_, elevators and
protractors attached to the metaphragma and base of the postfurca;
_z_, second flexor of hind leg; _a_, _a_, extensors of hind leg;
_c_, _c_, dorsal recti of abdomen. _Q_, ejaculatory duct; _R_,
penis; _S_, its prepuce. _M_, rectum.—After Straus-Durckheim, from
Newport.
]

Our knowledge of the muscular system of insects is still very
imperfect. To work it out thoroughly one should begin first with
that of Scolopendrella, then some generalized synapterous form, as
Japyx or Lepisma, then passing to that of a caterpillar, and ending
with some of the more highly specialized forms, such as a beetle,
etc. Thus far our knowledge is confined to that of the caterpillars
(Lyonet, Newport, and Lubbock) and the beetle (Straus-Durckheim) and
ants (Forel, Lubbock, and Janet).

=Musculature of a caterpillar.=—Newport’s account of that of the
larva of _Sphinx ligustri_ is the most useful (Fig. 231). The
muscles here present, he says, great uniformity of size and
distribution in every segment, the motions of each of these
divisions of the body being almost precisely similar, especially in
the 4th to 9th trunk segments. In these segments the first layer
seen on removing the fat and viscera are the flat straight recti
muscles. They are the most powerful of all the trunk muscles, and
are those which are most concerned in shortening the body, in
effecting the duplicature of the external teguments during the
changes of the insect, and which during the larval state mainly
assist in locomotion. There are four sets, two dorsal and two
ventral (Fig. 231, _A_, _A_). Without entering into farther details,
the reader is referred to the works of Newport and to Fig. 231.

=Musculature of a beetle.=—The best general account of the
musculature of a perfect insect is that of Straus-Durckheim in his
famous work on the Melolontha. We will copy the summary of Newport,
who adopted the nomenclature applied to these parts by Burmeister:—

“The muscles that connect the head with the thorax are contained
within the prothorax (Fig. 232, 2), and are of three kinds,
extensors, flexors, and retractors. The extensors, _levatores
capitis_ (_a_, _a_), consist of two pairs, one of which arises from
the middle line of the pronotum, and diverging laterally from its
fellow of the opposite side, passes directly forwards, and is
inserted by a narrow tendon into the anterior superior margin of the
occipital foramen. The other arises further back from the
prophragma. It is a long, narrow muscle that passes directly
forwards through the prothorax, and is inserted by a tendon near the
superior median line of the foramen; so that, while this muscle and
its fellow of the opposite side elevate the head almost in a
straight line, the one first described, when acting alone or singly,
draws the head a little on one side; but when the whole of these
muscles act in unison, they simply elevate the head upon the
prothorax. The depressors or flexors, _depressores capitis_ (_b_),
are exceedingly short muscles, which arise from the jugular plate,
or, when that part does not exist, from the border of the
prosternum, and are attached to the inferior margin of the occipital
foramen. They simply flex the head on the prothorax. The lateral
flexors, _depressores externi_ (_d_), are two little muscles that
arise from the same point as the preceding, and are attached to the
lateral inferior margin of the occipital foramen. The rotatory
muscles, _rotatores capitis_ (_c_), are two flat muscles like the
elevators, which arise, one at the side of the antefurca and the
other from the posterior jugular plate, and passing upwards and
outwards are attached to the lateral margin of the occipital
foramen. The _retractor_ or flexor of the jugular plate is a small
muscle (_e_) that arises from the margin of the antefurca, and
passing directly forwards is inserted by a small tendon into the
middle of the jugular piece. The _oblique extensor_ of the jugular
plate is a long, slender muscle (_f_) that arises from the external
margin of the pronotum, and passing obliquely downwards and forwards
traverses the prothorax and is inserted by a narrow tendon to the
jugular plate immediately before the retractor. The other retractor
(_g_) arises from the anterior superior boundary of the pronotum,
and passing downwards is inserted into the jugular plate between the
larger levator and _flexor capitis_.

“The muscles proper to the prothorax consist of four pairs, by which
it is united to the succeeding segments. The first of these, the
superior retractor, _retractor prothoracis superior_ (_h_), arises
by a broad, fleshy head from the anterior external margin of the
pronotum, and passing directly backwards is inserted by a tendon
into the prophragma, a little on one side of the median line. The
next muscle of importance, the inferior retractor (_i_), arises from
the anterior border of the medifurca, and is united to the posterior
of the antefurca, thus forming with that muscle part of the great
recti of the larva. This muscle must be considered as the proper
depressor of the prothorax. The _elevator prothoracis_ (_k_) is
narrow, pyramidal, and arises fleshy from the lateral surface of the
prophragma. It passes downwards and is attached by a narrow tendon
to the superior portion of the antefurca. The _rotatores
prothoracis_ are the largest of all the muscles of this segment.
They arise, one on each side (_l_), by a narrow head from the
posterior part of the pronotum, and passing beneath the prophragma
are considerably enlarged and attached to the tegument between the
two segments, and also to the anterior portion of the mesothorax.
The remaining muscle proper to the prothorax is the closer of the
spiracle, an exceedingly small muscle not shown in the drawing.

“The other muscles of this segment are those of the legs, which are
of considerable size. There are three distinct flexors of the coxa
(_m_, _n_, _o_). The first of these arises from the superior lateral
border of the pronotum, the second from the superior posterior
border, the third from the sides of the prothorax, and the fourth a
little nearer posteriorly, and the whole of them are attached by
narrow tendons to the sides of the coxa. But there is only one
extensor muscle to this part. In like manner, the extensor of the
trochanter is formed of three portions (Fig. 233, _a_, _b_, _c_);
but there is only one flexor (_d_), and one abductor (_e_). In the
femur, there is one extensor (_f_),—a long penniform muscle that
occupies the superior part of the thigh, and is attached by a tendon
to the anterior-posterior margin of the joint formed by the end of
the tibia. There is also but one flexor (_g_) in the femur, which,
like the preceding muscle, is penniform, and occupies the inferior
portion of the femur, and its tendon is attached to the inferior
border of the tibia. In the tibia itself there is also one flexor
and one extensor. The _flexor_ (_i_) occupies the superior portion
of the limb, and ends in a long tendon (_l_) that passes directly
through the joints of the tarsus, on their inferior surface, and is
attached to the inferior margin of the claw (_g_). The _extensor_
(_h_) occupies the inferior portion of the tibia, and is shorter
than the preceding muscle, like which it ends in a long tendon that
is attached to the upper margin of the claw. Besides these muscles,
which are common to the joints of the tarsus, there are two others
belonging to the claw, situated in the last joint. The first of
these, the _extensor_ (_m_), is short, and occupies the superior
portion of the last phalanx of the tarsus, and the other, the
_flexor_ (_n_), is a much longer penniform muscle, which occupies
nearly the whole of the upper and under surface of the posterior
part of the phalanx, and is attached, like the long flexor of the
tarsus, to the inferior part of the claw.”

[Illustration:

FIG. 233.—Muscles of the fore leg of _Melolontha vulgaris_: _a_,
_b_, _c_, three divisions of the extensor of the trochanter; _d_,
flexor,—_e_, abductor, of the trochanter; _f_, extensor of the
femur; _g_, flexor of the femur; _h_, extensor of the tibia; _i_,
flexor of the tibia; _l_, tendon attached to the lower edge of the
claw (_g_); _m_, extensor,—_n_, flexor, of the claw.—After
Straus-Durckheim, from Newport.
]

These are the muscles of the prothorax, and its organs of
locomotion. The reader is referred for a further account of the
muscles of the hinder thoracic and of the abdominal segments to
Straus-Durckheim’s original work.

=Minute structure of the muscles.=—The muscular fibres of insects are
striated (Figs. 235–238), even those of the alimentary canal; the only
notable exception being the alary muscles of the pericardial septum,
while Lowne states that certain of the thoracic muscles of the blow-fly
are not striated (Miall and Denny).

[Illustration:

FIG. 234.-Section through the prothorax of _Diapheromera femoratum_:
_prov_, proventriculus; _tr_, trachea; _n. c_, nervous cord; _s.
gl_, salivary gland; _hyp_, hypodermis; _ur. t_, urinary tube; _ht_,
heart; _m_, _m″_, _m‴_, muscles for lowering and raising the tergum;
_m′_, another muscle, its use unknown.
]

[Illustration:

FIG. 235.—Striated muscular fibre of Hydrophilus: _A_ and _B_, two
fibrillæ in a state of extension; _a_, thick disk; _b_, thin disk;
_c_, intermediate space. _C_, _D_, portion of the same fibrillæ seen
by moving the objective farther away and using a small diaphragm;
_n_, thick; _c_, thin disk. × 2000 diam.—After Ranvier, from
Perrier. _E_ after Gehuchten, from Lang.
]

In describing the minute structure of the muscles of ants, wasps, and
bees, C. Janet states that each consists of a group of fibres diverging
from a tendon, which is an integumentary invagination (Fig. 236). Each
fibre may be regarded as a multinucleate cell; the sarcolemma represents
the cell-membrane. It forms a resistant and extremely elastic tube. The
longitudinal (Fig. 236, _E_) and radiating filaments or reticulum
(spongioplasm of Gehuchten) lie in a nutritive filling substance (the
hyaloplasm of Gehuchten). The radiating filaments are formed of an
exceedingly elastic substance, and serve to sustain the longitudinal
filaments, to transmit the nervous stimulus to them, and to bring them
back into position after contraction. Janet’s account agrees on the
whole with that of Gehuchten.

[Illustration:

FIG. 236.—Preparations from the adductor muscle of the mandible of
_Vespa crabro_, worker, fixed by heat and alcohol several hours
after leaving its cell. _A_ to _E_ × 425; _F_ × 212: _A_, terminal
cupule of the tendon of a fibre. _B_, _C_, union of the fibres with
their tendon. _D_, branch of the tendon of a muscle sending out
tendons of some of the fibres; this branch is accompanied with
numerous nervous ramifications (_N_). _E_, fragment of a nerve which
furnishes the ramifications of Fig. _D_. _F_, fragment of the tendon
of the adductor muscle of the mandible; at the left are seen the
terminal cupules of the fibres (_td, c_); on the right, on the body
of the tendons, some sessile cupules, each of which forms the
attachment of a fibre; _td, b_, tendons of the fibres.—After Janet.
]

The muscles of flight are said to be penetrated by fine tracheal
branches, probably to supply a greater amount of oxygen, as the most
energetic movements of the insect are made in moving the wings during
flight; while the other muscles of the body are only surrounded by the
air-tubes. (Sharp.)

Without entering into tedious details, the reader is referred to figures
or references to the more important systems of muscles, such as those of
the legs and other appendages, of the wings, of respiration, etc., to
the sections treating of those organs or functions; also to Figs. 16,
17, 18, 22, 48, 74, 81, 83, 84, 115, 116, 172, 173, 174, etc.

=Muscular power of insects.=—The most detailed and careful experiments
are those of Plateau. His experiments prove that even the weakest
insects pull at least five times their own weight; many of them,
however, get the better of a burden twelve to twenty fold as heavy as
themselves, while a strong man or a draught horse, for example, is not
even able to pull a burden which is equal to the weight of his body.
Plateau came to the following results as to the relation of the weight
of the body to the load drawn (1 and 2 are to be compared with each
other, 1 being the larger, and 2 the smaller insect; it will be seen
that the smaller insect is the stronger).

[Illustration:

FIG. 237.—_Vespa crabro_, worker, fixed by heat and alcohol some hours
after leaving its cell. _A_ × 425; _B_ to _D_ × 850 times: _A_,
muscular fibre of the motor muscles of the mandibles treated, for
ten minutes, by 1 per cent potassium to bring out the reticulum; the
nodes of union of the rayed filaments with the longitudinal
filaments are indicated by distinct granulations (_l.d_), and these
longitudinal filaments present accessory thickenings (_d.a_); T,
trachea; N, junction of a nervous filament with the muscular fibres.
_B_, fibre of the same muscle, not treated with potassium, stained
by hæmatoxylin; _C_, transverse section of a disk at the level of a
layer of rayed filaments; _Sarc_, sarcolemma. _D_, transverse
section of a disk at the level of the rods; _nuc_, nucleus.—After
Janet.
]

1. _Carabus auratus_ 17.4.
2. _Nebria brevicollis_ 25.3.
1. _Cetonia aurata_ 15.
2. _Trichius fasciatus_ 41.3.
1. _Melolontha vulgaris_ 14.3.
2. _Anomala frischii_ 24.3.
1. _Oryctes nasicornis_ 4.7.
2. _Geotrupes stercorarius_ 9.8.
3. _Onthophagus nuchicornis_ 14.4.
1. _Necrophorus vespillo_ 15.1.
2. _Silpha livida_ 24.4.
1. _Ocypus morio_ 17.
2. _Quedeus fulgidus_ 29.6.
1. _Donacia nymphææ_, 42.7.
2. _Crioceris merdigera_ 39.2.
1. _Bombus terrestris_ 16.1.
2. _Bombus rupestris_ 14.5.
3. _Apis mellifica_ 20.2.

As regards the pushing power, the relation of the load to the size of
the body in different large beetles, gave the following figures:—

_Oryctes nasicornis_ 3.2.
_Geotrupes stercorarius_ 28.4.
_Onthophagus nuchicornis_ 92.9.

The leaping force of locusts was found by Straus-Dürckheim to be in
_Œdipoda grossa_ as 1.6, in _Œ. parallela_ as 3.3 of their weight.

[Illustration:

FIG. 238.—_Vespa crabro_, fixed and stained as in the subjects of the
other figures. _I_, _N_, _P_ × 1700; _H_, _J_, _M_ × 850; the others
× 425 times: _A-C_, motor muscles of the antennal scape. _D-P_,
motor muscles of the 3d coxa. _A_, _B_, the two ends, in very
different states of contraction, of the same fibre; on one side the
transverse striæ are near together, on the other very far apart.
_C_, a crushed and split fibre showing a fibrous appearance, owing
to the rupture of the radiated filaments, and the separation of the
longitudinal filaments. _D_, muscular disk seen in section, with two
rows of nuclei. _E_, a muscular fibre with three rows of nuclei.
_F_, a nucleus, accompanied with coagulated protoplasm, oozing from
a previous break of the muscular fibre. _G_, nerve-terminations very
near each other on the same muscular fibre. _H_, longitudinal
filaments, evenly covered with the coagulated substance, and
forming, throughout the mass of the fibre, continuous filaments.
_I_, filaments widely separated. _J_, longitudinal filaments showing
the beginning of one of the transverse breaks which isolate some of
the disks. _K_, oblique view of a disk obtained by such a break, and
of a fibre in circular section, with an axial row of nuclei; this
piece comprises three stages of radiated filaments. _L_, muscular
fibre with a row of nuclei; at the lower part, the nuclei have
issued from a longitudinal fissure in the fibre, and have remained
attached in a chain. _M_, edge of fibre in which there is quite a
large, clear space between the sarcolemma and the rods. _N_, passage
of the trachea, with the spiral thread, into three capillaries with
a smooth cuticula. _O_, elliptical disk from a fibre, with two rows
of nuclei, and showing a layer of radiated filaments. _P_, fragment
(highly magnified) of the edge of a disk seen in section.—After
Janet.
]

A humble bee (_Bombus terrestris_) can carry while flying a load 0.63 of
its own weight, and a honey bee 0.78; here, as usual, the smaller insect
is the stronger.[39]

LITERATURE ON THE MUSCLES

_a_. General

=Lyonet, P.= Traité anatomique de la chenille. La Haye, 1762.

=Cornalia, E.= Monographia del Bombyce del gelso. (Mem. R. Instituto
Lombardo Sc. Lett. ed Arte, 1856.)

=Basch, S.= Skelett und Muskeln des Kopfes von Termes. (Zeitschr. f.
wissens. Zool., xv, 1865, pp. 55–75, 1 Taf.)

=Lubbock, John.= Arrangement of the cutaneous muscles of the larva of
_Pygæra bucephala_. London, 1858. 2 Pls.

—— On some points in the anatomy of ants. (Month. Micr. Journ., xviii,
pp. 121–142, 1877, 4 Pls.)

—— On the anatomy of ants. (Trans. Linn. Soc., Ser. 2; Zool., ii,
1879, pp. 141–154, 2 Pls.)

=Poletajeff, N.= Du développement des muscles d’ailes chez les
Odonates. (Horæ Soc. Ent. Ross., xvi, 1879, pp. 10–37, 5 Pls.)

—— Die Flugmuskeln der Lepidopteren und Libelluliden. (Zool. Anzeiger,
1880, pp. 212, 213.)

—— Ueber die Flugmuskeln der Rhopaloceren. (Arbeiten d. Russ. Ent.
Ges., 1881, xiii, p. 9, 1 Taf., in Russian.)

=Lendenfeld, R. von.= Der Flug der Libellen. (Sitzb. k. Akad.
Wissens., 1 Abth. Wien, 1881, lxxxiii, pp. 289–376, 7 Taf.)

=Luks, Constantine.= Ueber die Brustmuskulature der Insekten. (Jena.
Zeitschr. f. Naturwissen., xvi, N. Folge IX, 1883, pp. 520–552, 2
Taf.)

=Carlet, G.= Sur les muscles de l’abdomen de l’abeille. (Comptes
rend., 1884, xcviii, pp. 758, 759.)

=Janet, Charles.= Sur les muscles des fourmis, des guêpes et des
abeilles. (Comptes rend., cxxi, p. 610, 1 Fig., 1895.)

Also the writings of Straus-Durckheim, Newport, Graber, Burgess,
Leydig, Dahl, Ockler, Dogiel, Dimmock, Kraepelin, Becher, Langer,
Kolbe.

_b_. Histology

=Aubert, H.= Ueber die eigenthümliche Struktur der Thoraxmuskeln der
Insekten. (Zeitschr. f. wissens. Zool., iv, 1853, pp. 388–399, 1
Taf.)

=Verson, E.= Zur Insertionsweise der Muskeln. (Sitzsb. Akad. d. wiss.
math. naturw. Cl. Wien., lvii, 1 Abth., pp. 63–66, 1868.)

=Künckel d’Herculais.= Sur le développement des fibres musculaires
striées chez les insectes. (Compt. rend. de l’Acad. Sc. Paris, lxxv,
1872.)

=Grunmach, Emil.= Ueber die Structur der quergestreiften Muskelfaser
bei den Insekten. Berlin, 1872. pp. 47.

=Fredericq, L.= Note sur la contraction des muscles striés de
l’Hydrophile. (Bull. Acad. Roy. Belgique, xli, p. 583, 2 Pls.)

=Gehuchten, A. van.= Étude sur la structure intime de la cellule
musculaire striée. (La Cellule, ii, pp. 289, 293–453, 1886, 6 Pls.)

=Janet, Charles.= Études sur les fourmis, les guêpes et les abeilles.
12^e note. (Structure des membranes articulaires des tendons et des
muscles, Limoges, 1895, pp. 25, 11 Figs.)

Also the writings of Burmeister, Chabrier, Leydig, Meckel, Lebert,
Wagner, Wagener, Amici, Krause, Heppner, Retzius, Rollet, G. Elias
Müller, F. Merkel, Hensen, Kölliker, Dogiel, Dönitz, Hagen,
Vosseler, Bütschli u. Schewiakoff, Lowne, Ciaccio, Biedermann,
Cohnbeim, Brücke, Haycraft, Melland, Bowman.

_c_. Muscular power of insects

=Plateau, Félix.= Sur la force musculaire des insectes. (Bull. Acad.
Roy. Belgique, 2 Sér. xx, 1865, pp. 732–757; xxii, 1866, pp.
283–308.)

—— Recherches sur la force absolue des muscles des invertébrés. 1884.

=Radan, R.= La force musculaire des insectes. (Revue de deux mondes, 2
Sér., lxiv, 1866, pp. 770–777.)

=Bibiakoff, Paul von.= Zur Muskelkraft der Insekten. (Natur, xvii,
1868, p. 399.)

=Delbœuf.= Nains et géants, Étude comparative de la force des petits
et des grands animaux. Bruxelles. (Also in Kosmos, xiii, 1883, pp.
58–62.)

=Camerano.= Mem. Acc. Torino (2), xliii, 1893, p. 229.

Also Newport, Art. Insecta, p. 76. Kirby and Spence, Burmeister,
Graber, Kolbe, pp. 375, 376.

THE NERVOUS SYSTEM

_a._ The nervous system as a whole

[Illustration:

FIG. 239.—Central nervous system of _Machilis maritima_: _au_, eye;
_lo_, optic tract; _g_, brain; _an_, antennal nerve; _oe_, œsophagus
passing between the œsophageal commissures; _usg_, infraœsophageal
ganglion; I-III, thoracic ganglia; 1–8, abdominal ganglia, the last
(_Sabc_) consisting of three fused ganglia; _s_, sympathetic nervous
system of the ventral cord.—After Oudemans, from Lang.
]

The nervous system of insects consists of a double series or chain of
ganglia connected by nervous cords or commissures. The first of these is
the brain or supraœsophageal ganglion; it is situated in the upper part
of the head, above the gullet or œsophagus, while the rest of the
system, called the ventral cord, lies on the floor of the body, under
the digestive canal.

A ganglion or nerve-centre consists of a mass of ganglion-cells, from
each of which a process or fibre passes off, uniting with others to form
a nerve; by means of these nerves the ganglia are connected with other
ganglia, and with the sensory cells and muscle-fibres. The ganglia may
be simple, and arranged in pairs, corresponding to each segment of the
body, or they may be compound, the result of the fusion of several pairs
of ganglia, which in the early stages of the embryo are separate. Thus
the brain of insects is a compound ganglion, or ganglionic mass.

The nerves are of two kinds: 1. _Sensory_, which transmit sensations
from the peripheral sense-cells to the ganglion, or brain; 2. _Motor_,
which send stimuli from the brain or any other ganglion to the muscles.

Of ganglion cells, some are tactile, and others give rise to nerves of
special sense, being distributed to the eyes, or to the organs of
hearing, smell, taste, or touch.

[Illustration:

FIG. 240.—Nervous system of _Melanoplus spretus_: _sp_,
supraœsophageal ganglion, sending off the large optic nerve (_op_)
to the eyes, and an ocellar nerve to each ocellus (the dotted line
_oc_ stops short of the left ocellus); _if_, infraœsophageal
ganglion; 1, 2, 3, thoracic ganglia; 1–5, five abdominal ganglia
(the fifth the largest, and sending branches to the ovipositor,
etc.) The sympathetic nerve and ganglia are represented by the two
main nerves which arise from the medio-cephalic (_as_) resting on
and above the œsophagus, and two ganglia (_ps_) on the under side of
the crop. From each of these ganglia, two nerves are sent under the
crop, and a larger nerve on each side to as far as the stomachal
cæca, ending the figure at the dotted line 2, near the second
thoracic ganglion. _u_, a round, shining body, connected by a nerve
with the medio-cephalic ganglion, its nature unknown.
]

[Illustration:

FIG. 241.—Section through the head of Machilis, showing the brain
(_br_), and subœsophageal ganglion (_soe. g_); _cl_, clypeus; _lbr_,
labrum; _oc_, ocellus.
]

While the supraœsophageal ganglion, or “brain,” of the insect is much
more complex than any other ganglion, consisting more exclusively both
of sensory as well as motor ganglia and their nerves, it should be borne
in mind that the subœsophageal ganglion also receives nerves of special
sense, situated on the palpi and on the tongue, as in the bee and other
insects; hence this ganglion is probably complex, consisting of sensory
and motor cells. The third thoracic ganglion is also, without doubt, a
complex one, as in the locusts the auditory nerves pass into it from the
ears, which are situated at the base of the abdomen, while in the green
grasshoppers, such as the katydids and their allies, whose ears are
situated in their fore legs, the first thoracic ganglion is a complex
one. In the cockroach and in Leptis (Chrysopila), a common fly, the
caudal appendages bear what are probably olfactory organs, and as these
parts are undoubtedly supplied from the last abdominal ganglion, this is
probably composed of sensory and motor ganglia; so that we have in the
ganglionated cord of insects a series of brains, as it were, running
from head to tail, and thus in a still stronger sense than in
vertebrates the entire nervous system, and not the brain alone, is the
organ of the mind of insects.

The simplest, most primitive form of the nervous system of insects is
seen in that of the Thysanura. That of Campodea has not yet been fully
examined, but in that of the more complicated genus, Machilis (Fig.
239), we see that there is a pair of ganglia to nearly each segment,
while the brain (Fig. 241) is composed of three lobes, viz. the optic,
the cerebral (Fig. 239, _g_), behind which is the antennal lobe, from
which the antennal nerve takes its origin. Behind the opening for the
throat (_oe_) is situated the first ganglion of the ventral cord, the
subœsophageal ganglion, which gives rise to the nerves supplying the
jaws and other mouth-parts.

[Illustration:

FIG. 242, _A-D._—The nervous systems of 4 genera of Diptera, to
demonstrate their various degrees of fusion of ganglia: _A_,
non-concentrated more primitive nervous system of _Chironomus
plumosus_, with 3 thoracic and 6 abdominal ganglionic masses. _B_,
nervous system of _Empis stercorea_, with 2 thoracic and 5 abdominal
ganglionic masses. _C_, nervous system of _Tabanus bovinus_, with 1
thoracic ganglionic mass, and the abdominal ganglia closely
approximated. _D_, highly modified nervous system of _Sarcophaga
carnaria_, in which all the ganglia of the ventral cord behind the
subœsophageal ganglion are fused into a single ganglionic
mass.—After Brandt, from Lang.
]

In the Collembola, which are retrograde Thysanura, there are from one
(Smynthurus), to three or four ventral ganglia.

In the winged insects, where the ganglia are more or less fused, the
fusion taking place in the head and at the end of the abdomen; there are
in the more simple and generalized forms, such as Ephemera, the
grasshopper, locusts (Fig. 240), etc., thirteen ganglia besides the two
pairs of compound ganglia in the head, three pairs of thoracic ganglia,
and usually from five to eight pairs of ganglia in the abdomen.

[Illustration:

FIG. 243.—Nervous system of the May beetle, _Lachnosterna fusca_:
_w^1_, nerve to 1st,—_w^2_, nerve to 2d, pair of wings; _ig_,
infraœsophageal ganglion.
]

[Illustration:

FIG. 244.—The same of the stag-beetle, _Lucanus dama_, where there are
3 thoracic, and 3 separate abdominal ganglia.
]

In certain winged insects the process of fusion or degeneration is
carried to such an extreme that there are either no abdominal ganglia
(Fig. 242, _D_), or their vestiges are situated in the thorax and
partially fused with the thoracic ones, as in the May beetle, in which
the prothoracic pair of ganglia is separate, while the two other
thoracic ganglia are fused with the abdominal, the latter being situated
in the thorax; this fusion is carried to a further extent than in any
other Coleoptera yet examined. In many Diptera and Hemiptera the
abdominal ganglia are either absent or the vestiges are fused with the
thoracic ganglia.

Rhizotrogus, which is allied to our May beetle, as also Hydrometra and
the Stylopidæ are said to lack the subœsophageal ganglion (Brandt).

In numerous Coleoptera (Acilius, Gyrinus, Necrophorus, Melolontha,
Bostrichus, Rhynchænus); in many Diptera (Culex, Tipula, Asilus,
Xylophaga, and Phora); and in the higher Hymenoptera (Crabronidæ,
Vespidæ, and Apidæ), as well as in many Lepidoptera (Vanessa, Argynnis,
and Pontia), two of the thoracic ganglia are fused together, while all
three are partially fused into a single mass in many brachycerous
Diptera (Conops, Syrphus, Pangonia, and the Muscidæ); in certain
Hemiptera (Pentatoma, Nepa, and Acanthia); also in a beetle (_Serica
brunnea_). Sometimes the subœsophageal ganglion is fused with the first
thoracic, as in Acanthia, Nepa, and Notonecta. The greatest amount of
variation is seen in the number of abdominal ganglia, all being fused
into a single one or from one to eight. The fusion is usually greatest
where the abdomen is shortened, due to the partial atrophy and
modification of the terminal segments which bear the ovipositor, where
present, and the genital armature.

There is only one pair of abdominal ganglia in Gyrinus and in certain
flies (Conops, Trypeta, Ortalis, and Phora); two in Rhynchænus, a
weevil, and in the flies, Syrphus and Volucella; three in Crabro and
Eucera; four in Sargus, Stratiomys and in butterflies, five in the
beetle, Silpha, and in the fly, Sciara, and the moth, Hepialus.

The nervous system in the larvæ of the metabolous orders is not
concentrated, though in that of the neuropterous Myrmeleo it has
undergone fusion from adaptation to the short compressed form of this
insect.

_b._ The brain

The brain of insects appears to be nearly, if not quite, as complex as
that of the lower vertebrates. As in the latter, the pair of
supraœsophageal ganglia, or brain, is the principal seat of the senses,
the chief organ of the insect’s mind.

It is composed of a larger number of pairs of primitive ganglia than any
of the succeeding nerve-centres, and is, structurally, entirely
different from and far more complicated than the other ganglia of the
nervous system. It possesses a central body in each hemisphere, a
“mushroom body,” optic lobes and optic ganglia and olfactory lobe, with
their connecting and commissural nerve-fibres, and a number of other
parts not found in the other ganglia.

In the succeeding ganglia the lobes are in general motor; the fibres
composing the œsophageal commissures, and which arise from the
œsophageal commissural lobes, extend not only to the subœsophageal
ganglion, but pass along through the succeeding ganglia to the last pair
of abdominal nerve-centres.[40] Since, then, there is a direct
continuity in the fibres forming the two main longitudinal commissures
of the nervous cord, and which originate in the brain, it seems to
follow that the movements of the body are in large part directed or
coördinated by the brain.[41] Still, however, a second brain, so to
speak, is found in the third thoracic ganglion of the locust, which
receives the auditory nerves from the ears situated in the base of the
abdomen; or in the first thoracic ganglion of the green grasshoppers
(katydids, etc.), whose ears are situated in their fore legs; while even
the last pair of abdominal ganglia in the cockroach and mole cricket,
is, so to speak, a secondary brain, since it distributes sensory nerves
to the caudal stylets, which are provided with organs probably olfactory
in nature.

It is impossible to understand the morphology of the brain unless we
examine the mode of origin of the nervous system in the early life of
the embryo. The head of an embryo insect consists of six segments,
_i.e._ the ocular, antennal, premandibular, mandibular, and the 1st and
2d maxillary segments, so named from the appendages they bear. Of these
the first three in the larva and adult are preoral, and the last three
are postoral. The antennal segment was probably either postoral in the
progenitors of insects, or the antennæ were inserted on the side of the
mouth, the latter finally moving back.[42]

The nervous system in the early embryonic condition, as shown by Wheeler
(Fig. 245), at first consists of nineteen pairs of primitive ganglia,
called _neuromeres_. Those of the head, which later in embryonic life
fuse together to form the brain, are the first three, corresponding to
the _protocerebrum_, _deutocerebrum_, and _tritocerebrum_ of Viallanes.
The first pair of primitive ganglia, and which is situated in front of
the mouth, is divided into three lobes.

[Illustration:

FIG. 245, _A-D_.—Diagrams of four consecutive stages in the
development of the brain and nerve-chain of the embryo of Xiphidium:
I, cephalic,—II, thoracic,—III, abdominal, region; _st_, stomodæum
or primitive mouth; _an_, anus; _e_, optic plate; _pc(og)_, 1st
protocerebral lobe, or optic ganglion; _pc^2_, _pc^3_, 2d and 3d
protocerebral lobes; _dc_, deutocerebrum; _tc_, tritocerebrum; 1–16,
the 16 postoral ganglia; _po. c_, postoral commissure; _fp_, furcal
pit; _ac_, anterior,—_pc_, posterior, ganglionic commissure; _ag_,
anterior,—_pg_, posterior,—_cg_, central,—_lg_, lateral
gangliomeres.—After Wheeler.
]

The first or outermost lobe, according to Wheeler, forms the optic
ganglion of the larva and imago, while the second and third lobes.
(_pc^2_, _pc^3_) ultimately form the bulk of the brain proper, or the
protocerebral lobes. The second (primitively postoral) brain-segment or
pair of ganglia gives origin to the antennæ, while the third brain, or
premandibular (intercalary) segment, gives origin to a temporary
embryonic pair of appendages found in Anurida and Campodea (the
premandibular ganglia), and also to the nerves supplying the labrum.
These three pairs of ganglia later on in embryonic life become preoral,
the mouth moving backwards. The three pairs of primitive ganglia,
behind, _i.e._ the mandibular and 1st and 2d maxillary ganglia, become
fused together to form the subœsophageal ganglion, and which in larval
and adult life is postoral.

If the tongue (ligula, or hypopharynx) represents a distinct pair of
appendages, then there are seven segments in the head.

[Illustration:

FIG. 246.—Section through head of a carabid, _Anopthalmus telkampfii_:
_br_, brain; _fg_, frontal ganglion; _soe_, subœsophageal ganglion;
_co_, commissure; _n_. _l_, nerve sending branches to the lingua
(_l_); _mn_, maxillary nerve; _mx_, 1st maxilla; _mm_, maxillary
muscle; _mx′_ 2d maxilla; _mt_, muscle of mentum; _le_, elevator
muscle of the œsophagus; _l′_ of the clypeus, and a third beyond
raising the labrum (_lbr_); _eph_, epipharynx; _g. g′_, salivary
glands above; _g^2_, lingual gland below the œsophagus (_œ_); _m_,
mouth; _pv_, proventriculus; _md_, mandible.
]

The brain, then, supplies nerves to the compound and simple eyes, and to
the antennæ, and gives origin to the sympathetic nerves; it is thus the
seat of the senses, also of the insect’s mind, and coördinates the
general movements of the body.

[Illustration:

FIG. 247.—Median longitudinal section through the head of _Blatta
orientalis_. The nervous system of the head is drawn entire. _hyp_,
hypopharynx; _os_. oral cavity; _lbr_, upper lip; _gf_, frontal
ganglion; _g_, brain; _na_, root of the antennal nerve; _no_, root
of the optic nerve; _ga_, anterior,—_gp_, posterior ganglion of the
paired visceral nervous system; _œ_, œsophagus; _c_, œsophageal
commissure; _usg_, infraœsophageal ganglia; _cc_, longitudinal
commissure between this and the first thoracic ganglion; _sg_,
common duct of the salivary glands; _lb_, labium (2d maxillæ); _nr_,
recurrent nerve; _d_, nerve uniting the frontal ganglion with the
œsophageal commissure; _e_, nerve from this commissure to the
labrum; _f_, nerve from the infraœsophageal ganglion to the
mandible, —_g_, to the 1st maxillæ, —_h_, to the lower lip (2d
maxillæ).—After Hofer, from Lang.
]

[Illustration:

FIG. 248.—1, front view of the brain of _Melanoplus femur-rubrum_:
_opt. gang_, optic ganglion; _oc_, ocelli and nerves leading to them
from the two hemispheres, each ocellar nerve arising from the region
containing the calices; _m. oc_, median ocellar nerve; _opt. l_,
optic lobe sending off the optic nerve to the optic ganglion; _ant.
l_, antennal or olfactory lobe; _ant. n_, antennal nerve; _f. g_,
frontal ganglion of sympathetic nerve; _lbr. n_, nerve to labrum;
_x_, cross-nerve or commissure between the two hemispheres; _œ. c_,
œsophageal commissure to subœsophageal ganglion. 2, side view of the
brain and subœsophageal ganglion (lettering of brain as in 1): _s.
g_, stomatogastric or sympathetic nerve; _a. s. g_, anterior, and
_p. s. g_, posterior, sympathetic ganglia; _g^2_, subœsophageal
ganglion; _md_, nerve to mandible; _mx_, maxillary nerve; _ln_,
labial nerve; _nl_, unknown nerve,—perhaps salivary. 3, interior
view of the right half of the head, showing the brain in its natural
position: _an_, antenna; _cl_, clypeus; _lbr_, labrum; _m_,
mouth-cavity; _md_, mandible; _t_, tongue; _œ_, œsophagus; _c_,
crop; _en_, right half of the endocranium or =X=-shaped bone,
through the anterior angle of which the œsophagus passes, while the
great mandibular muscles play in the lateral angles. The moon-shaped
edge is that made by the knife passing through the centre of the
=X=. 4, view of brain from above (letters as before). 5,
subœsophageal ganglion from above: _t. c_, commissure to the
succeeding thoracic ganglion (other letters as before). Fig. 3 is
enlarged 8 times; all the rest 25 times.—Drawn from original
dissections, by Mr. Edward Burgess, for the Second Report of the U.
S. Entomological Commission.
]

The pair of subœsophageal ganglia distributes nerves to the mandibles,
to the 1st and 2d maxillæ, and to the salivary glands (Fig. 248).

Its general shape and relations to the walls and to the outer organs of
the head is seen in Figs. 247, 248. In all the winged insects
(Pterygota) its plane is situated more or less at right angles to the
horizontal plane of the ventral cord. On the dorsal and anterior sides
are situated the ocular lobes, and below these the antennal lobes.

Viallanes first, independently of embryonic data, divided the brain of
adult insects into three regions or segments; _i.e._ the
“_protocerebron_,” “_deutocerebron_” and “_tritocerebron_,” which he
afterwards found to correspond with the three primitive elements
(neuromeres) of the brain and with the segments of the head of the
embryo.

The brain of the locusts (Melanoplus and Œdipoda) being best known will
serve as the basis of the following description, taken mainly from
Viallanes, with minor changes in the name of the three segments, and
other modifications.

=I.= =The optic or procerebral segment= is composed of a median portion,
_i.e._ two fused procerebral lobes (median protocerebrum), and of two
lateral masses, the optic ganglia (_protocerebrum_), and comprises the
following regions fused together and forming the median procerebral mass
(Viallanes):—

1. Procerebral lobes.
2. Optic ganglia.
3. Layer of postretinal fibres.
4. Ganglionic plate. (_Periopticon_ of Hickson.)
5. External chiasma.
6. External medullary mass. (_Epiopticon_ of Hickson.)
7. Internal chiasma.
8. Internal medullary mass. (_Opticon_ of Hickson.)
9. Optic ganglia and nerves.
10. Pedunculated or stalked body. (Mushroom body of Dujardin.)
11. Bridge of the procerebral lobes.
12. Central body.

[Illustration:

FIG. 249.—Diagram of an insect’s brain: _cc_, central body; _cg_,
ganglionic cells; _che_, external, _chi_, internal chiasma; _cœ_,
œsophageal commissure; _cp_, mushroom body; _ctc_, tritocerebral
commissure; _fpr_, postretinal fibres; _goc_, ocellar ganglion;
_goc_^1, œsophageal ganglion, the dotted ring the œsophagus; _gv^1_,
_gc^2_, _gv^3_, 1st, 2d, 3d, unpaired visceral ganglion; _gvl_,
lateral visceral ganglion; _ld_, dorsal lobe of the deutocerebrum;
_lg_, ganglionic plate; _lo_, olfactory lobe; _lpc_, protocerebral
lobe; _me_, external, _mi_, internal medullary mass; _na_, olfactory
or antennal nerve; _nl_, nerve to labrum; _no_, ocular nerve; _nt_,
tegumentary nerve; _œ_, œsophagus; _plp_, bridge of the
protocerebral lobes; _rvd_, visceral root arising from the
deutocerebrum; _rvt_, visceral root arising from the tritocerebrum;
_tr_, tritocerebrum; _to_, optic nerve or tract.—After Viallanes.
]

=Optic ganglia.=—Each of the two optic ganglia is formed of a series of
three ganglionic masses situated between the compound eyes and the
median procerebral mass, _i.e._ the ganglionic plate (Fig. 249, _lg_),
the external medullary mass (_me_), and the internal medullary mass
(_mi_).

The postretinal fibres (_fpr_) arising from the facets or single eyes of
the compound eye (ommatidia) pass into the ganglionic plate (_lg_),
which is united within by the chiasmatic fibres (_che_, external
chiasma) of the external medullary mass (_me_). The last is attached to
the internal medullary mass (_mi_) by fibres (_chi_), some of which are
chiasmatic, and others direct. Finally, the internal medullary mass
connects with the median part of the protocerebrum by direct fibres
forming the optic nerve or tract (_to_).

=Procerebral lobes.=—The median procerebral lobes are fused together on
the median line, forming a single central mass. From each side or lobe
arises the mushroom or stalked body. In the middle of the mass is the
central body, and directly in front is the procerebral bridge (_plp_).
The latter is a band uniting the two halves of the brain.

The procerebral lobes also give origin to the nerves to the ocelli
(_no_).

[Illustration:

FIG. 250.—Transverse section through the brain of the locust (Œdipoda
and Caloptenus): _c′_, lower part of the wall of _c_, calyx;—_st_,
stalk of the same; _bpcl_, bridge of the protocerebral lobes; _mo_,
nerve of median ocellus; _ch_, transverse fascia of the
optico-olfactory chiasma; _fcb_, fibrous region of the central body;
_lcb_, tubercle of the central body; _fch_, descending fascia of the
optico-olfactory chiasma; _choo_, superior fascia of the
optico-olfactory chiasma; _pt_, protocerebral lobes; _ld_, dorsal
lobe of the deutocerebrum; _lt_, tritocerebral lobe; _gcld_, _gc_,
ganglion cells.—After Viallanes.
]

=The mushroom or stalked bodies.=—These remarkable organs were first
discovered by Dujardin, who compared them to mushrooms, and observed
that they were more highly developed in ants, wasps, and bees than in
the lower insects, and thus inferred that the higher intelligence of
these insects was in direct relation to the development of these bodies.
We will call them the _mushroom bodies_.

These two bodies consist of a rounded lobular mass (the trabecula) of
the procerebral lobe, from which arises a double stalk (Fig. 253), the
larger called the _cauliculus_, the smaller the _peduncle_ (or pedicel);
these support the cap or _calyx_. The calices of the bee were compared
by Dujardin to a pair of disks on each side of the brain as seen from
above, “each disk being folded together and bent downwards before and
behind, its border being thickened, and the inner portion radiated.” In
the locust there are but two divisions of the calyx; in the cockroach,
ants, wasps, and bees, four.

The shape and relation of the mushroom bodies are represented in Figs.
252 and 253. The bodies are connected by commissural fibres, and are
connected with the optic ganglion of the same side, and with the central
body; while they are connected with the antennal lobes by the
optico-olfactory chiasma.

[Illustration:

FIG. 251.—Sagittal section through the brain of the locust: _l. oc.
n_, lateral ocellus nerve; _a. t_, anterior tubercle of the mushroom
body; _i. t_, internal tubercle of the mushroom body; _c. l_,
cerebral lobes; _l. l_, lateral lobe of the middle protocerebrum;
_com_, commissural cord; _c. mol_, central mass of the olfactory
lobe; _ac. an. l_, fibres uniting the median lobe of the middle
protocerebrum with dorsal lobes of the deutocerebrum; _gc. trit. l_,
ganglionated cortex of the tritocerebral lobe; _c. an. l_, cortex of
antennal (olfactory) lobe; _lab. fr_, labrofrontal nerve; _oe. com_,
œsophageal commissure; _tr. com_, transverse commissure of
œsophageal ring; other letters as in Fig. 250.—After Viallanes.
]

The stalked bodies are enveloped by the cortical layers of
ganglion-cells, those filling the hollow of the calyx having little or
no protoplasm around the nucleus.

=Structure of the mushroom bodies.=—By staining the brain of the
honey bee with bichromate of silver, Kenyon has worked out the
structure of the mushroom bodies, with their cells. The cup-shaped
bodies or calyces are composed of fibrillar substance
(_punktsubstanz_). Each of these cups, he says, is “filled to
overflowing with cells having large nuclei and very little
cytoplasm.” From the under surface of each of these cups there
descends into the general fibrillar substance of the brain “a column
of fibrillar substance, which unites with its fellow of the same
side to send a large branch obliquely downward to the median line of
the brain, and an equally large or larger branch straight forwards
to the anterior cerebral surface.”

The cells of the mushroom bodies, observes Kenyon, “stand out in
sharp contrast to all other nerve cells known, though they recall to
some extent the cells of Purkinje in the higher mammals. Each of the
cells contained within the fibrillar cup sends a nerve-process into
the latter, where it breaks up into a profusely arborescent system
of branchlets, which often appear with fine, short, lateral
processes, such as are characteristic of the dendrites of some
mammalian nerve-cells.” Just before entering the fibrillar
substance, a fine branch is given off that travels along the inner
surface of the cup along with others of the same nature, forming a
small bundle to the stalk of the mushroom body, down which it
continues until it reaches the origin of the anterior and the inner
roots above mentioned. “Here it branches, one branch continuing
straight on to the end of the anterior root, while the other passes
to the end of the inner root. Throughout its whole course the fibre
and its two branches are very fine. Nearly the whole stalk and
nearly the whole of each root is made up of these straight, parallel
fibres coming from the cells within the cup of the mushroom bodies.
What other fibres there are enter these bodies from the side, and
branch between the straight fibres very much as the dendrites of the
cells of Purkinje branch among the parallel fine fibres from the
cells of the granular layer in the mammalian cerebellum. These
fibres are of the nature of association fibres.”

Viallanes showed that from the olfactory or antennal lobes, as well
as from the optic ganglia, there are tracts of fibres which finally
enter the cups of the mushroom bodies, and Kenyon has confirmed this
observation. Kenyon has also, by the Golgi method, detected another
tract, before unknown, “passing down the hinder side of the brain,
from the cups to the region above the œsophagus, where it bends
forward and comes in contact with fibres from the ventral cord,
which exists, although Binet was unable to discover any growth of
fibres connecting the cord with the brain.

“The fibres entering the cups from the antennal lobe, the optic
ganglia, and the ventral region, spread out and branch among the
arborescent endings of the mushroom-body cells. The fibres branching
among the parallel fibres of the roots and the stalk lead off to
lower parts of the brain, connecting with efferent or motor-fibres,
or with secondary association fibres, that in their turn make such
connections. This portion of the circuit has not been perfectly made
out, though there seems to be sufficient data to warrant the
assumption just made.

[Illustration:

FIG. 252.—Section 17, showing the central body (_centr. b_) and
mushroom body, optic and antennal lobes (_a. l_), and procerebral
lobes (_pc. l_); _o. cal_, outer division of the calyx; _op. n_,
optic nerve; _trab_, trabeculum; _tc. n_, transverse nerve.
]

“Such fibres existing as described, there is then a complete circuit
for sensory stimuli from the various parts of the body to the cells
of the mushroom bodies. The dendritic or arborescent branches of
these cells take them up and pass them on out along the parallel
fibres or neurites in the roots of the mushroom bodies as motor or
other efferent impulses.

“This, however, is not all. For there are numerous fibres evident in
my preparations, the full courses of which I have not been thus far
able to determine, but which are so situated as to warrant the
inference that they may act as association fibres between the
afferent fibres from the antennæ, optic ganglia, and ventral system,
and the efferent fibres. There is then a possibility of a stimulus
entering the brain and passing out as a motor impulse without going
into the circuit of the fibres of the mushroom bodies; or, in other
words, a possibility of what may be compared to reflex action in
higher animals.”

[Illustration:

FIG. 253.—Enlarged view of the trabeculum (the dotted lines _tcn_ and
_obt. n_ pass through it) and its nerves, of the mushroom body,—its
calices and stalk, and the origin of the optic nerve × 225
diameters: _atn_, ascending trabecular nerve; _obt. n_, oblique
trabecular nerve; _tcn_, transverse nerve; _lat. n_, lateral nerve;
_cent. n_, central nerve.
]

The mushroom bodies have not yet been found to be present in the
Synaptera, but occur in the larvæ, at least of those of most metamorphic
insects (Lepidoptera and Hymenoptera), though not yet found in the larvæ
of Diptera. The writer has found these bodies in the nymphs of the
locust (_Melanoplus spretus_), but not in the embryo just before
hatching. They occur in the third larval or nymph stage of this insect.
It is evident that by the end of the first larval stage the brain
attains the development seen in the third larval state of the two-banded
species (_C. bivittatus_).

[Illustration:

FIG. 254.—Section through the brain of _Caloptenus bivittatus_ in the
third larval stage, showing the two hemispheres or sides of the
brain, and the ocelli and ocellar nerves, which are seen to arise
from the top of the hemispheres directly over the calices (compare
Fig. 251): _o. cal_, outer division of calyx of left mushroom body.
]

The result of our studies on the brain of the embryo locust was that
from the embryonic cerebral lobes are eventually developed the central
body and the two mushroom bodies. Fig. 254 shows the early condition of
the mushroom bodies and their undoubted origin from the cerebral
ganglia. Hence these bodies appear to be differentiations of the
cerebral ganglia or lobes, having no connection with the optic or
antennal lobes.

=The central body= (Fig. 252, _centr. b_).—This is the only single or
unpaired organ in the brain. Dietl characterizes it as a median
commissural system. Viallanes describes it as formed entirely of a very
fine and close fibrillar web, like a thick hemispherical skull-cap,
situated on the median line and united with the cerebral lobes. “It is
like a central post towards which converge fibres passing from all
points of the brain; being bound to the cerebral lobes, to the stalked
bodies, to the optic ganglia, and to the olfactory lobes by distinct
fibrous bundles.”

=The antennal or olfactory lobes (Deutocerebrum).=—This portion of the
brain consists of two hemispherical lobes, highly differentiated for
special sensorial perceptions, and connected by a slightly
differentiated medullary mass, the dorsal lobe (Figs. 248, 249 _lo_),
from which arise the motor fibres and those of general sensibility. The
antennal lobes are in part attached to the optic ganglia, and partly to
the stalked body on the same side, by the optic olfactory chiasma (Fig.
250 _fch_, _choo_), a system of fibres partially intercrossed on the
median line.

=The œsophageal lobes (Tritocerebrum)= (Figs. 249, 250).—From this
region the labrum and viscera are innervated, the nerves to the latter
being called the visceral, sympathetic, or stomatogastric system. As
Viallanes remarks, though plainly situated in front of the mouth, they
are in fact post-œsophageal centres. The two lobes are situated far
apart, and are connected by a bundle of fibres passing behind the
œsophagus, called the transverse commissure of the œsophageal ring
(Lienard). The œsophageal ganglia, besides giving rise to the labral
nerves, also give origin to the root of the frontal ganglion.

_c._ Histological elements of the brain

The brain and other ganglia are composed of two kinds of tissue.

1. The outer slightly darker, usually pale grayish white portion
consists of cortical or ganglion-cells differing in size. This portion
is stained red by carmine, the cells composing it readily taking the
stain.

The large ganglion cells (represented in Figs. 252 and 253) are oval,
and send off usually a single nerve-fibre; they have a thin fibrous
cell-wall, and the contents are finely granular. The nucleus is very
large, often one-half the diameter of the entire cell, and is composed
of large round refractive granules, usually concealing the nucleolus.

2. The medullary or inner part of the brain consists of matter which
remains white or unstained after the preparation has remained thoroughly
exposed to the action of the carmine. It consists of minute granules and
interlacing fibres. The latter often forms a fine irregular network
inclosing masses of finely granulated nerve matter.

This is called by Dietl “marksubstanz.” Leydig, in his Vom Bau des
thierischen Körpers, p. 89, thus refers to it:—

“In the brain and ventral ganglia of the leech, of insects, and
in the brain of the gastropods (Schnecken) I observe that the
stalks (stiele) of the ganglion-cells in nowise immediately
arise as nerve-fibres, but are planted in a molecular mass or
_punktsubstanz_, situated in the centre of the ganglion, and
merged with this substance. It follows, from what I have seen,
that there is no doubt that _the origin of the nerve-fibres
first takes place from this central punktsubstanz_.”

“This relation is the rule. But there also occur in the
nerve-centres of the invertebrates single, definitely situated
ganglion-cells, whose continuations become nerve-fibres without the
intervention of a superadded _punktsubstanz_.” We may, with Kenyon,
call it the fibrillar substance.

Leydig subsequently (p. 91) further describes this fibrillar
substance, stating that the granules composing it form a reticulated
mass of fibrillæ, or, in other words, a tangled web of very fine
fibres:—

“We at present consider that by the passage of the continuation of
the ganglion-cells into the _punktsubstanz_ this continuation
becomes lost in the fine threads, and on the other side of the
_punktsubstanz_ the similar fibrillar substance forms the origin of
the axis-cylinders arranged parallel to one another; so it is quite
certain _that the single axis-cylinder derives its fibrillar
substance as a mixture from the most diverse ganglion-cells_.”

_d._ The visceral (sympathetic or stomatogastric) system

This system in insects is composed (1) of a series of three unpaired
ganglia (Fig. 249, _gv^1_, _gv^2_, _gv^3_), situated over the
dorso-median line of the œsophagus, and connected by a median nervous
cord or recurrent nerve (_nr_, _vagus_ of Newport). The first of these
ganglia is the frontal ganglion, which is connected with the œsophageal
ganglia by a pair of roots (_rvt_), which have an origin primitively
common with that of the labral nerves (Fig. 248, _fg_ and _lbr_).

[Illustration:

FIG. 255.—Anterior portion of the paired and unpaired visceral nervous
system of _Blatta orientalis_ seen from above. The outlines of the
brain (_g_) and the roots of the antennal nerve (_na_), which cover
a portion of the sympathetic nervous system, are given by dotted
lines. Lettering as in Fig. 247. _nsd_, nerve to salivary gland. The
nervus recurrens (_nr_) enters an unpaired stomach ganglion farther
back.—After Hofer, from Lang.
]

2. Of two pairs of lateral ganglia (Fig. 255, _ga_, _gp_) situated two
on each side of the œsophagus. They are connected both with the antennal
lobes by a nerve (_rvd_), and to the chain of unpaired ganglia by a
special connective. The first pair of these ganglia sends nerves to the
heart and aorta; the second pair to the tracheæ of the head.

The unpaired median or recurrent nerve (_nr_) extends back from under
the brain along the upper side of the œsophagus, and (in Blatta), behind
the origin of the nerves to the salivary glands, enters an unpaired
ganglion, called the stomachic ganglion (_ganglion ventriculare_),
situated in front of the proventriculus. The number of these stomachic
ganglia varies in different orders of insects.

In Blatta, Küpffer and also Hofer have shown (Fig. 255) (Müller,
Brandt, _ex_ Kolbe) that the nerve to each salivary gland arises
from three different centres: the anterior end situated under the
œsophagus is innervated by the paired visceral nerves from the
hinder paired ganglia; the remaining part by nerves arising from
each side of the recurrent nerve; and thirdly by a pair of nerves
arising from the subœsophageal ganglion which accompanies the common
salivary duct, and ends in branches which partly innervate the
salivary glands and in part their muscles.

Hofer considers that the function of this complex system of paired and
unpaired ganglia, with their nerves, is a double one, viz. serving both
as a centre for the peristaltic action of the œsophagus, and as
innervating the salivary glands.

Besides these a second portion of the visceral system arises from the
thoracic and abdominal ventral cord. It may be seen in the simplest
condition yet known in the nervous system of Machilis (Fig. 239 _s_). It
consists of a fine, slender nerve, which extends along the surface of
the ventral chain of ganglia, and sending off a pair of branches
(accessory transverse nerves) in front of each ganglion. These accessory
nerves receive nerve-twigs from the upper cord of the ventral chain,
dilating near their origins into a minute elongated ganglion, and then
passing partly outwards to the branches of the tracheæ and the muscles
of the spiracles, uniting in the middle line of each segment of the body
behind the head, _i.e._ of those segments containing a pair of ganglia.

_e._ The supraspinal cord

In the adult Lepidoptera has been detected, continuous with and on the
upper side of the abdominal portions of the ventral cord, a longitudinal
cord of connective tissue forming a white or yellowish band, and which
seems to be an outgrowth of the dorsal portion of the neurilemma of the
ventral cord. Muscles pass from it to the neighboring ventral portions
of the integument. Its use is unknown, and attention was first called to
it by Treviranus, who called it “an unknown ventral vessel”
(_Bauchgefäss_). Afterwards it was re-discovered by Newport, who
described it as “a distinct vascular canal.” But Burger has proved by
cross-sections that it is not tubular, but a comparatively solid cord
composed, however, of loose connective tissue. Newport found it in the
larva of _Sphinx ligustri_, but Cattie states that it is not present in
that of _Acherontia atropos_. It has not yet been observed in insects of
other orders, but its homologue exists in the scorpion and in the
centipede, and it may prove to correspond with the far more complete
arterial coat which, with the exception of the brain, envelops the
nervous system of Limulus.

_f._ Modifications of the brain in different orders of insects

There are different grades of cerebral development in insects, and
Viallanes claimed that it was no exaggeration to say that the brain of
the locust (Melanoplus) differs as much from that of the wasp as that of
the frog differs from that of man. He insists that the physiological
conditions which determine the anatomical modifications of the brain are
correlated with 1, the food; 2, the perfection of the senses; and 3,
with the perfection of the psychic faculties. For example, in those
which feed on solid food and whose œsophagus is large (Orthoptera and
Coleoptera), the connectives are elongated, the subœsophageal commissure
free in all its extent, and the tritocerebrum is situated quite far from
the preceding segment of the brain.

On the other hand, in insects which feed on fluid food (Hymenoptera,
Lepidoptera, Diptera, Hemiptera), the œsophagus is slender and the
nervous centres which surround them are very much condensed; the
connectives are short, and the tritocerebrum is closely fused, partly to
a portion of the antennal lobes (deutocerebrum) and partly to the
mandibular ganglion.

As regards the perfection of the senses, where, as in dragon-flies, the
eyes are very large, the optic ganglia are correspondingly so, and in
the same insects the antennæ being very small, the antennal lobes are
almost rudimentary. The ants exhibit inverse conditions; in their brain
the antennal lobes are well developed, while the optic ganglia are
reduced, and where, as in Typhlopone, the eyes are wanting, they are
completely atrophied.

[Illustration:

FIG. 256.—Head of _Anophthalmus tellkampfii_, showing the brain,—the
optic ganglia, nerves, and eyes totally atrophied.
]

[Illustration:

FIG. 257.—Head of another Carabid, with the brain and eyes normal:
_op_, optic ganglion; _pcl_, brain.
]

In certain cave insects where the eyes are wanting, the optic ganglia
are also absent. In the eyeless cave species of Anophthalmus the optic
ganglia and nerves are entirely atrophied, as they are in Adelops,
which, however, has vestiges of the facets (ommatidia). Fig. 257
represents the brain of _Chlænius pennsylvanicus_, a Carabid beetle,
with its eyes and optic ganglia (_op_) which may be compared with
Anopthalmus, in which these parts are totally atrophied.

Dujardin claimed that the degree of complication of the stalked body of
the Hymenoptera was in direct relation with their mental powers. This
has been proved by Forel, who has shown that in the honey bee and ants
the mushroom bodies are much more developed in the workers than in the
males or females and Viallanes adds that these bodies are almost
rudimentary in the dragon-flies, whose eyes are so large; while on the
contrary in the blind ants (Typhlopone), these bodies are as perfect and
voluminous as in the ants with eyes.

[Illustration:

FIG. 258.—Diagrammatic outlines of sections of the upper part of the
brain of a cockroach. Only one side of the brain is here
represented. The numbers indicate the position in the series of 34
sections into which this brain was cut. _mb_, mushroom bodies, with
their cellular covering (_c_) and their stems (_st_); _a_, anterior
nervous mass; _m_, median nervous mass.—After Newton.
]

Within the limits of the same order the stalked bodies are most perfect
in the most intelligent forms. Thus in the Orthoptera, says Viallanes,
the Blattæ, Forficulæ, and the crickets, the mushroom bodies are more
perfect than in the locusts, which have simpler herbivorous habits. This
perfection of the mushroom bodies is seen not only in the increase in
size, but also in the complication of its structures. Thus in the groups
with lower instincts (Tabanus, Æschna) the stalk does not end in a calyx
projecting from the surface of the brain, but its end, simply truncated,
is indicated externally only by an accumulation of the ganglionic nuclei
which cover it.[43]

In types which Viallanes regards as more advanced, _i.e._ Œdipoda and
Melanoplus, the end of the stalk projects and is folded into a calyx.

The brain of the cockroach (Periplaneta, Fig. 258) is a step higher than
that of the locusts, each calyx being divided into two adjacent calices,
although the cockroaches are an older and more generalized type than
locusts.

The stalked bodies of cockroaches are thus complex, like those of the
higher Hymenoptera, the calices in Xylocopa, Bombus, and Apis being
double and so large as to cover almost the entire surface of the brain.

Finally, in what Viallanes regards as the most perfect type (Vespa), the
sides of the calices are folded and become sinuous, so as to increase
the surface, thus assuming an appearance which, he claims, strongly
recalls that of the convolutions of the brain of the mammals.

Cheshire also calls attention to a progression in the size of these
appendages, as well as in mental powers as we rise from the
cockchafer (_Melolontha vulgaris_) to the cricket, up to the
ichneumon, then to the carpenter bee, and finally to the social hive
bee, “where the pedunculated bodies form the ⅕ part of the volume of
the cerebral mass, and the 1⁄870 of the volume of the entire
creature, while in the cockchafer they are less than 1⁄2300 the
part. The size of the brain is also a gauge of intelligence. In the
worker bee the brain is 1⁄174 of the body; in the red ant, 1⁄296; in
the Melolontha, 1⁄3500; in the Dyticus beetle, 1⁄4400.” (Bees and
bee-keeping, p. 54.)

_g._ Functions of the nerve-centres and nerves

As we have seen, the central seat of the functions of the nervous system
is not the brain alone (supraœsophageal ganglion), but each ganglion is
more or less the seat of vital movements, those of the abdomen being
each a distinct motor and respiratory centre. The two halves of a
ganglion are independent of each other.

According to Faivre, the brain is the seat of the will and of the power
of coördinating the movements of the body, while the infraœsophageal
ganglion is the seat of the motive power and also of the will.

The physiological experiments of Binet, which are in the line of those
of Faivre, but more thorough, demonstrate that an insect may live for
months without a brain, if the subœsophageal ganglion is left intact,
just as a vertebrate may exist without its cerebrum. As Kenyon says:
“Faivre long ago showed that the subœsophageal ganglion is the seat of
the power of coördination of the muscular movements of the body. Binet
has shown that the brain is the seat of the power directing these
movements. ‘A debrained hexapod will eat when food is placed beneath its
palpi, but it cannot go to its food even though the latter be but a very
small space removed from its course or position. Whether the insect
would be able to do so if the mushroom bodies only were destroyed, and
the antennal lobes, optic lobes, and the rest of the brain were left
intact, is a question that yet remains to be answered’” (Kenyon).

In insects which are beheaded, however readily they respond to
stimulation of the nerves, they are almost completely wanting in will
power. Yet insects which have been decapitated can still walk and fly.
Hymenoptera will live one or two days after decapitation, beetles from
one to three days, and moths (Agrotis) will show signs of life five days
after the loss of their head.

That the loss of will power is gradual was proved by decapitating
_Polistes pallipes_. A day after the operation she was standing on her
legs and opening and closing her wings; 41 hours after the operation she
was still alive, moving her legs, and thrusting out her sting when
irritated. _Ichneumon otiosus_, after the removal of its head, remained
very lively, and cleaned its wings and legs, the power of coördination
in its wings and legs remaining. A horse-fly, a day after decapitation,
was lively and flew about in a natural manner.[44]

When the abdomen is cut off, respiration in that region is not at first
interrupted. The seat of respiratory movements was referred by Faivre to
the hinder thoracic ganglion, but Plateau says that this view must be
entirely abandoned, remarking: “All carefully performed experiments on
the nervous system of Arthropoda have shown that each ganglion of the
ventral chain is a motor centre, and in insects a respiratory centre,
for the somite to which it belongs” (Miall and Denny’s The Cockroach, p.
164).

The last pair of abdominal ganglia serve as the nervous centre of the
nerves sent to the genital organs.

The recurrent or stomatogastric nerve, which, through the medium of the
frontal ganglion, regulates digestion, has only a slight degree of
sensibility; the insect remains quiet even when a powerful allurement is
presented to the digestive tract (Kolbe).

Faivre states that the destruction of the frontal ganglion, or a section
of the commissures connecting it with the brain, puts an end to
swallowing movements; on the other hand, stimulation results in
energetic movements of this nature.

Yersin, by cutting through the commissure in different places, and thus
isolating the ganglia of the nervous cord of _Gryllus campestris_,
arrived at the following results:—

1. The section of a nerve near its origin rendered the organ supplied by
this nerve incapable of performing its functions.

2. If the connectives between two ganglia, _i.e._ the second and third
thoracic ganglia, are cut through, the fore as well as hinder parts of
the body retain their power of motion and sensation; but a stimulus
applied to the anterior part of the body does not pass to the hinder
portion.

3. Insects with an incomplete metamorphosis after section of the
connectives are not in every case unable to moult and to farther
develop.

4. If only one of the two connectives be cut through, the appendages of
the side cut through which take their origin between the place injured
and the hinder end of the body, often lose sensation and freedom of
motion, or the power of coördination of movements becomes irregular.
Sometimes this is shown by an unsteadiness in the gait, so that the
insect walks around in a circle; after a while these irregularities
cease, and the movements of the limbs on the injured side are only
slightly restrained. By a section of both connectives in any one place
the power of coördination of movements is not injured.

5. The section of the connectives appear to have no influence on
nutrition, but affects reproduction, the attempt at fertilization on the
part of the male producing no result, and the impregnated female laying
no eggs.

6. Injury to the brain, or to the subœsophageal, or one of the thoracic
ganglia, is followed by a momentary enfeeblement of the ganglion
affected. Afterwards there results a convulsive trembling, which either
pervades the whole body or only the appendages innervated by the injured
ganglion.

7. As a result of an injury to the brain there is such a lack of
steadiness in the movements that the insect walks or flies in a circle;
for instance, a fly or dragon-fly thus injured in flying describes a
circle or spiral. Steiner, in making this experiment, observed that the
insect circled on its uninjured side. The brain is thus a motor centre.

8. By injuring a thoracic ganglion, one or all the organs which receive
nerves from the ganglion are momentarily weakened. Afterwards the
functions become restored. Sometimes, however, the insect walks in a
circle. Faivre observed that after the destruction of the metathoracic
ganglion of _Dyticus marginalis_ the hind wings and hind legs were
partially paralyzed (Kolbe, _ex_ Yersin).

LITERATURE ON THE NERVOUS SYSTEM

_a._ General

=Newport, George.= On the nervous system of the _Sphinx ligustri_ L.,
and on the changes which it undergoes during a part of the
metamorphoses of the insect. (Phil. Trans. Roy. Soc., London, 1832,
pp. 383–398; 1834, pp. 389–423, Pls.)

=Helmholtz, H. L. F.= De fabrica systematis nervosi evertebratorum.
Diss. in aug. Berolini, 1842.

=Blanchard, E.= Recherches anatomiques et zoologiques sur le système
nerveux des animaux sans vertèbres. Du système nerveux des insectes.
(Annales des Sciences nat., Sér. 3, v, 1846, pp. 273–379, 8 Pls.)

—— Du système nerveux chez les invertèbres dans ses rapports avec la
classification de ces animaux. Paris, 1849.

—— in Cuvier’s Règne animal. (Edition accompagnée de planches gravées.
Insectes. Pl. 3, 3_a_, and 4.)

=Leidy, Joseph.= History and anatomy of the hemipterous genus
Belostoma. (Memoirs Amer. Acad. Arts and Sc., N. S. iv, 1849, pp.
57–67, 1 Pl.)

=Scheiber, S. H.= Vergleichende Anatomie und Physiologie der
Œstridenlarven. (Sitzungsb. k. Akad. wiss. Wien. Math.-Naturwiss.
Cl., xli, 1860, pp. 439–496; xlv, 1862, pp. 7–68; 5 Taf.)

=Tullberg, Tycho.= Sveriges Podurider. (K. Svenska vet. Akad. Handl.
x, 1872, pp. 1–70, 12 Taf.)

=Berlese, A.= Osservazione sulla anatomia descrittiva del _Gryllus
campestris_ L. (Atti della soc. Veneto-Trentina, 1880, vii, pp.
200–299.)

=Baudelot, E.= Contributions à la physiologie der système nerveux des
insectes. (Revue d. sc. nat., i, pp. 269–280, 1872.)

=Studer, Th.= Ueber Nervenendigung bei Insekten. Kleine Beiträge zur
Histologie der Insekten. (Mitt. Naturf. Ges., Bern, 1874, pp.
97–104, 1 Taf.)

=Brandt, E.= Recherches anatomiques et morphologiques sur le système
nerveux des insectes Hyménoptères. (Compt. rendus de l’Acad. Sc.,
Paris, 1875.)

—— Ueber das Nervensystem der Apiden. (Sitzungsb. d. naturf. Ges., in
Petersbourg, vii, 1876.)

—— Ueber das Nervensystem der Schmetterlingsraupen. (Verhandl. der
Russ. Ent. Gesellsch., x, 1877. Also 16 other articles with plates,
in Horæ Soc. Ent. Ross., 1878–1882.)

=Mark, E. L.= The nervous system of Phylloxera. (Psyche, ii, pp.
201–207, 1879.)

=Riley, Charles Valentine.= The nervous system and salivary glands of
Phylloxera. (Psyche, ii, pp. 225, 226, 1879.)

=Cholodkowsky, N.= Zur Frage über den Baue und über die Innervation
der Speicheldrüsen der Blattiden. (Horæ Soc. Ent. Ross., 1881, xvi,
pp. 6–9, 2 Taf.)

=Liénard, V.= Constitution de l’anneau œsophagien. (Archives de
Biologie, i, pp. 381–391, 1880, 1 Taf.)

=Michaels, H.= Nervensystem von _Oryctes nasicornis_ im Larven-,
Puppen-, und Käferzustande. (Zeits. f. wissens. Zool., xxxiv, 1880,
pp. 641–702, 4 Taf.)

=Rossi, A.= Sul modo di terminare dei nervi nei muscoli dell’ organo
sonoro della Cicala commune (_Cicada plebeja_). (Mem. accad. sc.
Bologna, 1880, 4 Ser., i, pp. 661–665.)

=Foettinger, A.= Sur le termination des nerfs dans les muscles des
insectes. (Archiv de Biologie, i, 1880.)

=Binet.= Contribution à l’étude der system nerveux sous intestinal des
insectes. (Journ. l’anat. et phys., xxx, pp. 449–580, 1894.)

=Paulowa.= Zum Bau des Eingeweide Nervensystems der Insekten. (Zool.
Anzeiger., xviii, Feb. 25, 1895, pp. 85–87.)

Also the writings of Lyonet, Cuvier, Rolando, Straus-Durckheim,
Leydig, Newport, Graber, Viallanes, Grassi, Oudemans.

_b._ The brain

=Dujardin, F.= Mémoires sur le système nerveux des insectes. (Annales
des Sciences nat, Sér. 3, 1850, xiv, pp. 195–206, Pl. 1, 1850.)

=Rabl-Rückhard.= Studien über Insectengehirne. (Archiv für Anatomie,
Physiologie, etc., herausg. von Reichert u. R. du Bois-Raymond,
1876, p. 480, Taf. i.)

=Dietl, M. J.= Die Organization des Arthropodengehirns. (Zeitschr.
wissens. Zool., xxvii, 1876, p. 488, Taf. xxxvi.-xxxviii.)

=Flogel, T. H. L.= Ueber den einheitlichen Bau des Gehirns in den
verschiedenen Insectenordnungen. (Zeitschr. wissens. Zool., xxx,
Suppl., 1878, p. 556, Taf. xxiii, xxiv.)

=Newton, E. T.= On a new method of constructing models of the brains
of insects, etc. (Journ. Quekett Microscopical Club, pp. 150–158,
1879.)

—— On the brain of the cockroach, _Blatta orientalis_. (Quart. Journ.
Microscopical Science, July, 1879, p. 340, Pl. xv, xvi.)

=Packard, A. S.= The brain of the locust. (Chapter xi, Second Report
of the U. S. Entomological Commission, pp. 223–242, Pls. ix-xv,
1880.)

=Cuccati, Giovanni.= Sulla stuttura del ganglio sopraesofageo di
alcuni ortotteri. (Acrydium lineola, Locusta viridissima, Locusta
(species?), Gryllotalpa vulgaris, Bologna, 1887, 4º, pp. 1–27, Pl.
i-iv.)

—— Intorno alla struttura del cervello della Sonomya erythrocephala,
nota preventiva. Bologna, 1887.

—— Ueber die Organization des Gehirns des Sonomya erythrocephala.
(Zeitschr. f. wissens. Zool., 1888, xlvi, pp. 240–269, 2 Taf.)

=Viallanes, H.= Études histologiques et organologiques sur les centres
nerveux et les organes des sens des animaux articulés.

1. Mémoire. Le ganglion optique de la langouste (_Palinurus
vulgaris_). (Annal. d. Sc. Nat. Zool., 1884, 6^e Sér., xvii, Art. 3,
pp. 1–74, 5 Pls.)

2. Mémoire. Le ganglion optique de la Libellule (_Æschna
maculatissima_). (Ibid., 1885, 6^e Sér., xviii, Art. 4, pp. 1–34, 3
Pls.)

3. Mémoire. Le ganglion optique de quelques larves de Diptères
(_Musca_, _Eristalis_, _Stratiomys_). (Ibid., 1886, 6^e Sér., xix,
Art. M. 4, pp. 34, 2 Pls.)

4. Mémoire. Le cerveau de la guêpe (_Vespa crabro et vulgaris_).
(Ibid., 1887, 7^e Sér., ii, pp. 5–100, 6 Pls.)

5. Mémoire. 1. Le cerveau du criquet (_Œdipoda cœrulescens_ et
_Caloptenus italicus_). 2. Comparaison du cerveau des Crustacés et
des Insectes. 3. Le cerveau et la morphologie du squelette
céphalique. (Ibid., 1888, 7^e Sér., iv, pp. 1–120, 6 Pls.)

—— Sur la structure interne du ganglion optique de quelques larves de
Diptères. (Bull. Soc. Phil., Paris, 1885, 7^e Sér., ix, pp. 75–78.)

—— La structure du cerveau des Hyménoptères. (Bull. Soc. Philomat.,
Paris, 1886, 7^e Sér., x, pp. 82, 83.)

—— La structure du cerveau des Orthoptères. (Bull. Soc. Philomat.,
Paris, 1886, 7^e Sér., xi, pp. 119–126.)

—— Sur la morphologie comparée du cerveau des Insectes et des Crustacés.
(Compt. rend. Acad. Sc. Paris, 1887, civ, pp. 444–447.)

=Kenyon, F. C.= The meaning and structure of the so-called “mushroom
bodies” of the hexapod brain. (Amer. Naturalist, xxx, 1896, pp. 643–650,
1 fig.)

—— The brain of the bee. (Journ. Comp. Neurology, vi, fasc. 3, 1896, pp.
133–210.)

—— The optic lobes of the bee’s brain in the light of recent
neurological methods. (Amer. Nat., xxxi, 1897, pp. 369–376, 1 Pl.)

With the embryological works of Graber, Heider, Korscheldt, Patten,
Wheeler, etc.

_c._ Histology of the nervous System

=Helmholtz.= De fabrica systematis nervosi evertebratorum. Diss.
Berolini, 1842.

=Remak.= Ueber d. Inhalt d. Nervenprimitivröhren. (Archiv f. Anat. u.
Phys., 1843.)

=Leydig.= Lehrbuch der Histologie der Menschen und der Thiere. 1857.

—— Vom Bau des thierischen Körpers. i. 1864.

—— Tafeln zur vergleichenden Anatomie. i. Tübingen, 1864.

—— Zelle und Gewebe, neue Beiträge zur Histologie des Tier-Körpers.
Bonn, 1885, pp. 219, 6 Taf.

=Walter.= Mikroscopische Studien über das Centralnervensystem
wirbelloser Thiere. 1863.

=Dietl, M. J.= Die Gewebselemente des Centralnervensystems bei
wirbellosen Thieren. (Aus den Berichten des naturw.-medic. Vereins
in Innsbruck.) Innsbruck, 1878.

=Berger.= Untersuchungen über den Bau des Gehirns und der Retina der
Arthropoden. (Arbeiten des zool. Instituts zu Wien, Heft 2, p. 173,
1878.)

—— Nachtrag zu den Untersuchungen über den Bau des Gehirns und der
Retina der Arthropoden. (Ibid., Heft 3.)

=Viallanes, H.= Recherches sur l’histologie des insectes, etc. Paris,
1882. (Annales des Sciences nat., pp. 1–348, Pls. 1–18.)

—— Sur la structure de la substance ponctuée des insectes. Paris,
1885.

=Haller, B.= Ueber die sogenannte Leydig’sche Punktsubstantz im
Centralnervensystem. (Morp. Jahrb., xi, 1886.)

=Nansen, F.= The structure and combination of the histological
elements of the central nervous system. (Bergen’s Museum
Aarsberetning for 1886. Bergen, 1887.)

Also the writings of Benedicenti, Holmgren.

THE SENSORY ORGANS

_a._ The eyes and insect vision

[Illustration:

FIG. 259.—Different forms of compound eyes. _A_, a bug (Pyrrhocoris).
_B_, worker bee. _C_, drone. _D_, male Bibio, a holoptic
insect.—From Judeich and Nitsche.
]

Of the eyes of insects there are two kinds, the simple and the compound.
Of the former there are usually three, arranged in a triangle near the
top of the head, between the compound eyes (Fig. 259, _B_). The compound
or facetted eyes, which are usually round and prominent, differ much in
size and in the number of facets.

The number of facets varies from 12 in Lepisma,—though in a
Brazilian beetle (Lathridius) there are only seven unequal
facets,—to 50 in the ant, and up to 4000 in the house-fly, 12,000 in
_Acherontia atropos_, 17,000 in Papilio, 20,000 in the dragon-fly
(Æschna), 25,000 in a beetle (Mordella), while in _Sphinx
convolvuli_, the number reaches 27,000. The size of the facets seems
to bear some relation to that of the insect, but even in the
smallest species none have been observed less than 1⁄2000 of an inch
in diameter. Day-flying Lepidoptera have smaller facets than moths
(Lubbock).

[Illustration:

FIG. 260.—Section through the ocellus of a young Dyticus larva: _ct_,
cuticula; _l_, corneal lens; _gh_, cells of the vitreous body, being
modified hypodermal cells (_hy_); _st_, rods; _re_, retinal cells;
_no_, optic nerve.—After Grenacher, from Lang.
]

=The simple, or single-lensed eye (ocellus).=—Morphologically the simple
eye is a modified portion of the ectoderm, the pigment enclosing the
retinal cells arising from specialized hypodermal cells, and covered by
a specialized transparent portion of the cuticula, forming the corneal
lens. The apparatus is supplied with a nerve, the fibres of which end in
a rod or solid nerve-ending, as in other sensory organs.

As seen in the ocellus of Dyticus (Fig. 260), under the corneal lens the
hypodermis forms a sort of pit, and the cells are modified to form the
vitreous body (vitrella) and retina. Each retinal cell (_re_) is
connected with a fibre from the optic nerve, contains pigment, and ends
in a rod directed outwards towards the lens. The cells at the end of the
pit or depression are, next to the lens, without pigment, and, growing
in between the retina and the lens, fill it up, and thus form a sort of
vitreous body.

The ocellus appears to be a direct heirloom from the eyes of worms,
while the many-facetted compound eye of the crustaceans and of
insects is peculiar to these classes. The compound eye of the
myriopod Scutigera differs structurally in many respects from the
compound eye of insects, and that of Limulus still more so.

It should be observed that in the young nymph of Ephemera, as well
as in the semipupa of Bombus, each of the three ocelli are situated
on separate sclerites. In Bombus the anterior ocellus has a double
shape, being broad, transversely ovate, and not round like the two
others, as if resulting from the fusion of what were originally two
distinct ocelli.

The ocelli are not infrequently wanting, as in adult Dermaptera, in
the Locustidæ, and in certain Hemiptera (Hydrocora). In Lepidoptera
there are but two ocelli; in geometrid moths they are often
atrophied, and they are absent in butterflies (except Pamphila).

=The compound or facetted eye (ommateum).=—The facetted arthropod eye is
wonderfully complex and most delicately organized, being far more so
than that of vertebrates or molluscs. The simplest or most primitive
facetted eye appears to be that of Lepisma. As stated by Watase, the
compound eye of arthropods is morphologically “a collection of
ectodermic pits whose outer open ends face towards the sources of light,
and whose inner ends are connected with the central nervous system by
the optic nerve fibres.”

The facetted eye is composed of numerous simple eyes called _ommatidia_,
each of which is complicated in structure. The elements which make up an
_ommatidium_ are the following: (1) The facet or cornea, which is a
specialized portion of the cuticula; and (2), the crystalline lens or
cone; (3), the nerve-ending or _retinula_, which is formed out of the
retinula cells and the _rhabdom_ or rod lying in its axis; and (4) of
the pigment enclosing the lens and rod; the last three elements are
derived from the hypodermis. The single eyes are separated from each
other by pigment cells.

=The facet or cornea.=—This is biconvex, clear, transparent, usually
hexagonal in outline, and refracts the light. The corneal lenses are
cast in moulting.

The corneal lenses are circular in most cases where they are very
convex, as in Lathridius and Batocera. The hexagonal ones are very
irregular. When they are very convex the eye has a granular
appearance, but when not greater than the convexity of the eye
itself, the eye appears perfectly smooth (Bolbocerus, etc.). The
facets in the lower part of the eye of Dineutes are a trifle larger
than in the upper part (about nine to ten). In many insects the
reverse is the case, the upper facets being larger than the lower, a
notable instance being Anax. The intervening lines between the
facets are often beset with hairs, sometimes very long and dense, as
in the drone bee and Trichophthalmus; and the modifications of the
hairs into scales which takes place on the body occurs on the eyes
also, the scales on the eyes of some beetles of the family Colydiidæ
being very large, arranged in lines over the eyes like tombstones
(Trachypholis).[45]

[Illustration:

FIG. 261.—Section through the eye of a fly (_Musca vomitoria_): _c_,
cornea, or facet; _pc_, pseudocone; _r_, retinula; _Rh_, rhabdom;
_pg^1_, _pg^2_, _pg^3_, pigment cells; _b.m_, basilar membrane; _T_,
_Tt_{1}_, _Tt_{2}_, trachea; _tv_, tracheal vesicle; _t.a_, terminal
anastomosis; _op_, opticon; _c.op_, epiopticon; _p.op_, periopticon;
_n.c_, nuclei; _n.c.s_, nerve-cell sheath; _N.f_, decussating
nerve-fibres.—After Hickson, from Lubbock.
]

=The crystalline lens or cone.=—Behind or within the facets is a layer
composed of the cones, behind which are the layers of retinulæ and
rhabdoms, and which correspond to the layer of rods and cones, but not
the retina as a whole, of vertebrate animals.

The crystalline lens is, when present, usually more or less conical, and
consists of four or more hypodermis-cells.

The cones are of various shapes and sizes in insects of different
groups, or are entirely wanting, and Grenacher has divided the eyes of
insects into _eucone_, _pseudocone_, and _acone_. As the pseudocone
seems, however, to be rather a modification of the eucone eye, the
following division may be made:—

1. _Eucone eyes_, comprising those with a well-developed cone. They
occur in Lepisma, Blatta (Fig. 262), and other Orthoptera, in
Neuroptera, in Cicadidæ, in those Coleoptera with five tarsal joints, in
the dipterous genus Corethra, and in the Lepidoptera and Hymenoptera
(Fig. 263).

[Illustration:

FIG. 262.—Ommatidium of cockroach (Periplaneta): _lf_, cornea; _kk_,
crystalline cone; _pg′_ pigment cell; _rl_, retinula; _rm_,
rhabdom.—After Grenacher, from Lubbock.
]

[Illustration:

FIG. 263.—Two separate elements of the eucone eye of a bee; _Lf_,
cornea; _n_, nucleus of Semper; _Kk_, crystalline cone; _Pg_,
pigment cells; _Rl_, retinula; _Rm_, rhabdom.—After Grenacher, from
Lubbock.
]

[Illustration:

FIG. 264.—Three ommatidia of a pseudocone eye, diagrammatic: _A_, a
separate ommatidium of _Musca vomitoria_, semi-diagrammatic: _c_,
cornea; _p.c_, pseudocone; _pg′_, pigmented cells surrounding the
pseudocone; _p.g_{2}_, additional pigment cells; _p.g_{3}_, basal
pigment cells; _n.p.c_, nuclei of pseudocone; _r_, retinulæ; _n.r_,
_n.r′_, nucleus of retinulæ; _R_, rhabdom; _b.m_, basal membrane;
_t.a_, terminal anastomosis sending nerve-fibrils to the retinulæ.
_B_, section through a retinula and rhabdom near the basal membrane,
the six retinulæ (_r_) fused into a tube ensheathing the rhabdom
(_R_).—After Hickson.
]

_a._ Pseudocone eyes; in which, instead of the crystalline lens or cone,
there are four cells filled with a transparent fluid medium, and a
smaller protoplasmic portion containing a nucleus (Muscidæ, Fig. 264,
_pc_). Hickson states that the difference between the eucone and
pseudocone eyes lies in the fact that in the pseudocone eye “the
refracting body formed by the cone-cell lies behind the nuclei,” and in
the eucone eye in front of it.

2. _Acone eyes_, where the cone or refracting body is wanting, but is
represented by the four primitive cone-cells. Acone eyes occur in
Forficulidæ, Hemiptera (except Cicadidæ), the nematocerous Diptera
(Tipula, etc.), and those Coleoptera which have less than five tarsal
joints.

=The retinula and rod.=—The retinula is morphologically a nerve-end
cell, situated at the end of a nerve-fibril arising from the optic
nerve. The elements of the retinula of Musca are six in number and
surround the _rhabdom_ (Fig. 264), which consists of a bundle of six
long, delicate chitinous rods, more or less firmly united together (Fig.
264, _R_).

The six elements of the retinula of Musca are in their outer or distal
portion free from one another, but towards their base are fused into a
sheath (Fig. 264, _r_). They are true nerve-end cells, as shown by
Müller and by Max Schultze, their views having been confirmed by
Grenacher and by Hickson. The relations of the nerves to the rods after
passing through the basal membrane is seen in Fig. 266.

=The pigment.=—The cones or pseudocones are mostly buried in pigment, as
well as the rods; and the pigment forms two layers. The outer of the two
layers is called the iris pigment (Fig. 265, _e_, _iris tapetum_), and
the inner (_f_) the retinal pigment.

Between the ommatidia internally there occur, according to Hickson,
pigment cells (Fig. 264, _p.g_{3}_), each of which stands on the basilar
membrane and sends a fine process outwards towards the internal process
of the external pigment-cell (_p.g_{2}_). A long, slender tracheal
vesicle also passes in between the retinulæ.

[Illustration:

FIG. 265.—Two ommatidia from the eye of _Colymbetes fuscus_, × 160:
_a_, cornea; _b_, cone; _c_, rhabdom; _d_, basal membrane, with
nerve filaments below it: _e_, iris pigment; _f_, retina
pigment.—After Exner, from Sharp.
]

=The basilar membrane.=—This is a thin fenestrate membrane (Fig. 261)
separating the cones and rods from the optic tract (Fig. 264, _b.m_). It
is perforated for the passage of tracheal diverticula and of the optic
nerve fibrils. It separates the dioptric or instrumental portion of the
eye from the percipient portion, _i.e._ the optic tract.

=The optic tract.=—This is the optic ganglion of earlier writers, and
appears to be the percipient portion of the eye, as opposed to the
dioptric portion. If the reader will examine Figs. 249 and 261, he will
see that it consists of three distinct ganglionic swellings, _i.e._ the
_opticon_, _epiopticon_, and _periopticon_, whose structure is very
complicated. In Musca (Fig. 261) the first ganglionic swelling (opticon)
is separated from the brain by a slight constriction, which Berger
regards as the homologue of the optic nerve of the other arthropods. It
consists of a very fine granular matrix traversed throughout by a fine
meshwork of minute fibrillæ, the neurospongium of Hickson. In the young
cockroach (Periplaneta) the optic nerve separating the cerebral ganglion
from the opticon is much longer in proportion than it is in the adult
blow-fly.

[Illustration:

FIG. 266.—Periopticon and terminal anastomosis of Agrion, showing the
character of the elements of the periopticon (_p.op_) and the
structure of the terminal anastomosis (_t.a_). 1. The first layer of
the terminal anastomosis, consisting of a plexus of fibrils and
nerve-cells (_n.c_). 2. The second layer, in which the fibrils are
collected together in bundles. 3. The final optic plexus and
nerve-cells. 4. The layer in which the optic fibrils are collected
in bundles to be distributed to the retinulæ (_r_); _b.m_, basal
membrane.—After Hickson.
]

The second ganglionic swelling (epiopticon, Fig. 261, _c.op_) is
separated from the opticon by a tract of fine nerve-fibrils, which
partially decussate; at the decussation two or three larger nerve-cells
may be seen. It also contains a few scattered nerve-cells (_n.c_). The
third ganglionic swelling (periopticon, _p.op_) is separated from the
others by a bundle of long optic nerve-fibrils, which cross one another.
It is composed of a number of cylindrical masses of neurospongium
arranged side by side (Fig. 261, _p.op_). Between these elements of the
periopticon, which do not seem to bear any relation to the number of
ommatidia, a single nerve-cell is very frequently seen. The periopticon
does not occur in Periplaneta and Nepa (Hickson). The three optic
ganglia thus described, together with the cerebral ganglia, are
surrounded by a sheath of densely packed nerve-cells.

Bearing in mind the fact that the retinulæ are the nerve-end cells
of the fibres passing through the periopticon, it will be well to
read the following account, by Hickson, of the terminal anastomosis
of the optic fibrils in the periopticon of _Agrion bifurcatum_, and
to examine his sketch (Fig. 266):

“The terminal anastomosis of Agrion may be conveniently divided into
four regions. First the region (1) lying nearest to the periopticon
in which the nerve-cells are numerous, and the fibrils leaving the
periopticon form a complicated plexus; the region (2) next to this,
in which the fibrils have collected into bundles separated by spaces
occupied by very thin-walled tracheæ in which there are no spiral
markings, and lymph-spaces; next, the region (3) in which the
fibrils form a final plexus, and in which there are again a
considerable number of nerve-cells; and, lastly, the region (4) in
which the fibrils are again collected into bundles, separated by
spaces containing tracheæ, which perforate the basement membrane to
supply the retinulæ.”

It would seem as if the decussation of the optic nerve-fibrils were
a matter of primary importance, as it so generally occurs, but in
the young of that most generalized of all pterygote insects, the
cockroach (Periplaneta), Hickson states that the optic nerve-fibrils
which leave the periopticon pass without decussating to the
ommateum, and in the adult there is only a partial decussation. In
Nepa there is no decussation, but the anastomosis is complicated by
the presence of looped and transverse anastomoses.

Looking at the eye as a whole, Hickson regards all the nerve structure
of the eye lying between the crystalline cone-layer and the true optic
nerve to be analogous with the retina of other animals. With Ciaccio,
Berger, and others, he does not regard the layer composed of the
retinulæ and rhabdoms as the equivalent of the retina of vertebrates,
etc.

=Origin of the facetted eye.=—The two kinds of eye, the simple and the
compound, are supposed to have been derived from a primitive type,
resembling the single eye (ommatidium) of the acone eye of Tipula. As
stated by Lang, “an increase of the elements of this primitive eye led
to the formation of the ocellus; an increase in number of the primitive
eyes, and their approximation, led to the formation of the compound
facet eye.” This view is suggested, he says, by the groups of closely
contiguous single eyes of the myriopods, considered in connection with
the compound eye of Scutigera. Grenacher looks upon simple (ocelli) and
compound eyes as “sisters,” not derived from one another, but from a
common parentage.

Immature insects rarely possess compound eyes; they are only known
to occur in the nymphs of Odonata and Ephemeridæ, and in the larvæ
and pupa of Corethra.

=Mode of vision by single eyes or ocelli.=—In their simplest condition,
the eyes of worms and other of the lower invertebrates, probably only
enable those animals to distinguish light from darkness. The ocelli of
spiders and of many insects, however, probably enable them, as Lubbock
remarks, to see as our eyes do. The simple lens throws on the retina an
image, which is perceived by the fine terminations of the optic nerve.
The ocelli of different arthropods differ, however, very much in degree
of complexity.

Müller considered that the power of vision of ocelli “is probably
confined to the perception of very near objects.”

“This may be inferred,” Müller states, “partly from their existing
principally in larvæ and apterous insects, and partly from several
observations which I have made relative to the position of these
simple eyes. In the genus Empusa the head is so prolonged over the
middle inferior eye that, in the locomotion of the animal, the
nearest objects can only come within the range. In _Locusta
cornuta_, also, the same eye lies beneath the prolongation of the
head.... In the Orthoptera generally, also, the simple eyes are, in
consequence of the depressed position of the head, directed
downwards towards the surface upon which the insects are
moving.”[46] Lowne considers that in the ocellus of Eristalis, the
great convexity of the lens must give it a very short focus, and the
comparatively small number of rods render the picture of even very
near objects quite imperfect and practically useless for purposes of
vision, and that the function of the ocelli is “the perception of
the intensity and the direction of light, rather than of vision, in
the ordinary acceptation of the term.”

Réaumur, Marcel de Serres, Dugès, and Forel have shown by
experiment, that in insects which possess both ocelli and compound
eyes, the former may be covered over without materially affecting
the movements of the animals, while if the facetted eyes are
covered, they act as if in the dark (Lubbock).

While Plateau regards the ocelli as of scarcely any use to the
insect, and Forel claims that wasps, humble bees, ants, etc., walk
or fly almost equally well without as with the aid of their ocelli,
Lubbock demurs to this view, and says the same experiments of
Forel’s might almost be quoted to prove the same with reference to
the compound eyes. Indeed, the writer has observed that in caves,
eyeless beetles apparently run about as freely and with as much
purpose, as their eyed relatives in the open air.

Plateau has recently shown that caterpillars which have ocelli alone
are very short-sighted, not seeing objects at a distance beyond one
or two centimetres, and it has been fully proved by Plateau and
others, that spiders, with their well-formed ocelli, are myopic, and
have little power of making out distinctly the shape of the objects
they see.

On the whole, we are rather inclined to agree with Lubbock and
Forel, that the ocelli are useful in dark places and for near
vision. They are, as Lubbock states, especially developed in
insects, such as ants, bees, and wasps, which live partly in the
open light and partly in the dark recesses of nests. Moreover, the
night-flying moths nearly all possess ocelli, while with one known
exception (Pamphila) they are wanting in butterflies.

Finally, remarks Lubbock, “Whatever the special function of ocelli
may be, it seems clear that they must see in the same manner as our
eyes do—that is to say, the image must be reversed. On the other
hand, in the case of compound eyes, it seems probable that the
vision is direct, and the difficulty of accounting for the existence
in the same animal of two such different kinds of eyes is certainly
enhanced by the fact that, as it would seem, the image given by the
medial eyes is reversed, while that of the lateral ones is direct”
(p. 181).

=Mode of vision by facetted eyes.=—The complexity of the facetted eyes
of insects is amazing, and difficult to account for unless we accept the
mosaic theory of Müller, who maintained that the distinctness of the
image formed by such an eye will be greater in proportion to the number
of separate cones. His famous theory is thus stated: “An image formed by
several thousand separate points, of which each corresponds to a
distinct field of vision in the external world, will resemble a piece of
mosaic work, and a better idea cannot be conceived of the image of
external objects which will be depicted on the retina of beings endowed
with such organs of vision, than by comparing it with perfect work of
that kind.”

[Illustration:

FIG. 267.—From Lubbock.
]

How vision is effected by a many-facetted eye is thus explained by
Lubbock: “Let a number of transparent tubes, or cones with opaque walls,
be ranged side by side in front of the retina, and separated from one
another by black pigment. In this case the only light which can reach
the optic nerve will be that which falls on any given tube in the
direction of its axis.” For instance, in Fig. 267, the light from _a_
will pass to _a′_, that from _b_ to _b′_, that from _c_ to _c′_, and so
on. The light from _c_, which falls on the other tubes, will not reach
the nerve, but will impinge on the sides and be absorbed by the pigment.
Thus, though the light from _c_ will illuminate the whole surface of the
eye, it will only affect the nerve at _c′_.

According to this view those rays of light only which pass directly
through the crystalline cones, or are reflected from their sides, can
reach the corresponding nerve-fibres. The others fall on, and are
absorbed by, the pigment which separates the different facets. Hence
each cone receives light only from a very small portion of the field of
vision, and the rays so received are collected into one spot of light.

It follows from this theory that the larger and more convex the eye, the
wider will be its field of vision, while the smaller and more numerous
are the facets, the more distinct will be the vision (Lubbock).

The theory is certainly supported by the shape and size and the immense
number of facets of the eye of the dragon-fly, which all concede to see
better, and at a longer range, than probably any other insect.

Müller’s mosaic theory was generally received, until doubted and
criticised by Gottsche (1852), Dor (1861), Plateau, and others. As
Lubbock in his excellent summary states, Gottsche’s observation
(previously made by Leeuwenhoek) that each separate cornea gives a
separate and distinct image, was made on the eye of the blow-fly,
which does not possess a true crystalline cone. Plateau’s objection
loses its force, since he seems to have had in his mind, as Lubbock
states, Gottsche’s, rather than Müller’s, theory.

Müller’s theory is supported by Boll, Grenacher, Lubbock, Watase,
and especially by Exner, who has given much attention to the subject
of the vision of insects, and is the weightiest authority on the
subject.

Gottsche’s view that each of the facetted eyes makes a distinct
image which partially overlaps and is combined with all the images
made by the other facets, was shown by Grenacher to be untenable,
after repeating Gottsche’s experiments with the eyes of moths, in
which the crystalline cones are firm and attached to the cornea. He
was thus able to remove the soft parts, and to look through the
cones and the cornea. When the microscope was focussed at the inner
end of the cone, a spot of light was visible, but no image. As the
object-glass was moved forward, the image gradually came into view,
and then disappeared again. Here, then, the image is formed in the
interior of the cone itself.

Exner attempted to make this experiment with the eye of Hydrophilus,
but in that insect the crystalline cones always came away from the
cornea. “He, however, calculated the focal length, refraction, etc.,
of the cornea, and concluded that, even if, in spite of the
crystalline cone, an image could be formed, it would fall much
behind the retinula.”

“In these cases, then,” adds Lubbock, “an image is out of the
question. Moreover, as the cone tapers to a point, there would, in
fact, be no room for an image, which must be received on an
appropriate surface. In many insect eyes, indeed, as in those of the
cockchafer, the crystalline cone is drawn out into a thread, which
expands again before reaching the retinula. Such an arrangement
seems fatal to any idea of an image.”

Lubbock thus sums up the reasons which seem to favor Müller’s theory
of mosaic vision, and to oppose Gottsche’s view: “(1) In certain
cases, as in Hyperia, there are no lenses, and consequently there
can be no image; (2) the image would generally be destroyed by the
crystalline cone; (3) in some cases it would seem that the image
would be formed completely behind the eye, while in others, again,
it would be too near the cornea; (4) a pointed retina seems
incompatible with a clear image; (5) any true projection of an image
would in certain species be precluded by the presence of
impenetrable pigment, which only leaves a minute central passage for
the light-rays; (6) even the clearest image would be useless, from
the absence of a suitable receptive surface, since both the small
number and mode of combination of the elements composing that
surface seem to preclude it from receiving more than a single
impression; (7) no system of accommodation has yet been discovered;
finally (8), a combination of many thousand relatively complete eyes
seems quite useless and incomprehensible.”

In his most recent work (1890) on the eyes of crustacea and insects,
Exner states that the numerous simple eyes which make up the
compound eye have each a cornea, but it is more or less flat, and
the crystalline part of the eye has not the shape of a lens, but of
a “lens cylinder,” that is, of a cylinder which is composed of
sheets of transparent tissue, the refracting powers of which
decrease toward the periphery of the cylinder. If an eye of this
kind is removed and freed of the pigment which surrounds it, objects
may be looked at through it from behind; but its field of vision is
very small, and the direct images received from each separate eye
are either produced close to one another on the retina (or rather
the retinulæ of all the eyes) or superposed. In this last case no
less than thirty separate images may be superposed, which is
supposed to be of great use to night-flying insects. Exner claims
that many other advantages result from the compound nature of an
insect’s eye. Thus the mobile pigment, which corresponds to our
iris, can take different positions, either between the separate eyes
or behind the lens cylinders, in which case it acts as so many
screens to intercept the over-abundance of light. Exner finds that
with its compound eyes the common glow-worm (Lampyris) is capable of
distinguishing large signboard letters at a distance of ten or more
feet, as well as extremely fine lines engraved one-hundredth of an
inch apart, if they are at a distance of less than half an inch from
the eye. Exner substantiates the truth of the results of Plateau’s
experiments, and claims that while the compound eye is inferior to
the vertebrate eye for making out the forms of objects, it is
superior to the latter in distinguishing the smallest movements of
objects in the total field of vision.

More recently Mallock has given some optical reasons to show that
Müller’s view is the true one. He concludes, and thus agrees with
Plateau, that insects do not see well, at any rate as regards their
power of defining distant objects, and their behavior certainly
favors this view. It might be asked, What advantage, then, have
insects with compound eyes over those with simple eyes? Mallock
answers, that the advantage over simple-eyed animals lies in the
fact that there is hardly any practical limit to the nearness of the
objects they can examine. “With the composite eye, indeed, the
closer the object the better the sight, for the greater will be the
number of lenses employed to produce the impression; whereas, in the
simple eye the focal length of the lens limits the distance at which
a distinct view can be obtained.” He gives a table containing
measures of the diameters and angles between the axes of the lenses
of various insect eyes, and states that the best of the eyes would
give a picture about as good as if executed in rather coarse
woodwork and viewed at a distance of a foot, “and although a distant
landscape could only be indifferently represented on such a
coarse-grained structure, it would do very well for things near
enough to occupy a considerable part of the field of view.”

=The principal use of the facetted eye to perceive the movements of
animals.=—Plateau adopts Exner’s views as to the use of the facetted eye
in perceiving the movements of other animals. He therefore concludes
that insects and other arthropods with compound eyes do not distinguish
the form of objects; but with Exner he believes that their vision
consists mainly in the perception of moving bodies.

Most animals seem but little impressed by the form of their enemies
or of their victims, though their attention is immediately excited
by the slightest displacement. Hunters, fishermen, and entomologists
have made in confirmation of this view numerous and demonstrative
observations.

Though the production of an image in the facetted eye of the insect
seems impossible, we can easily conceive, says Plateau, how it can
ascertain the existence of a movement. Indeed, if a luminous object
is placed before a compound eye, it will illuminate a whole group of
simple eyes or facets; moreover, the centre of this group will be
clearer than the rest. Every movement of the luminous body will
displace the centre of clearness; some of the facets not illuminated
will first receive the light, and others will reënter into the
shade; some nervous terminations will be excited anew, while those
which were so formerly will cease to be. Hence the facetted eyes are
not complete visual organs, but mainly organs of orientation.

Plateau experimented in the following way: In a darkened room, with
two differently shaped but nearly equal light-openings, one square
and open, the other subdivided into a number of small holes, and
therefore of more difficult egress, he observed the choices of
opening made by insects flying from the other end of the room.
Careful practical provisions were made to eliminate error; the
light-intensity of the two openings was as far as possible equalized
or else noted, and no trees or other external objects were in view.
The room was not darkened beyond the limit at which ordinary type
ceases to be readable, otherwise the insects refused to fly (it is
well known that during the passage of a thick cloud insects usually
cease to fly). These observations were made on insects both with or
without ocelli, in addition to the compound eyes, and with the same
results.

From repeated experiments on flies, bees, etc., butterflies and
moths, dragon-flies and beetles, Plateau concludes that insects with
compound eyes do not notice differences in form of openings in a
half-darkened room, but fly with equal readiness to the apparently
easy and apparently difficult way of escape; that they are attracted
to the more intensely lighted opening, or to one with apparently
greater surface; hence he concludes that they cannot distinguish the
form of objects, at least only to a very slight extent, though they
readily perceive objects in motion.

One result of his experiments is that insects only utilize their
eyes to choose between a _white_ luminous orifice in a dark chamber,
or another orifice, or group of orifices, _equally white_. They are
guided neither by odorous emanations nor by differences of color. He
thinks that bees have as bad sight and act almost exactly as flies.

From numerous experiments on Odonata, Coleoptera, Lepidoptera,
Diptera, and Hymenoptera Plateau arrives provisionally at the
following conclusions:

1. Diurnal insects have need of a quick strong light, and cannot
direct their movements in partial obscurity.

2. Insects with compound eyes do not notice differences of form
existing between two light orifices, and are deceived by an excess
of luminous intensity as well as by the apparent excess of surface.
In short, they do not distinguish the form of objects, or if they
do, distinguish them very badly.

Lubbock, however, does not fully accept Plateau’s experiments with
the windows, and thinks they discern the form of bodies better than
Plateau supposes.

=How far can insects see?=—It is now supposed that no insects can
perceive objects at a greater distance than about six feet. On an
average Lepidoptera can see the movements of rather large bodies
1.50 meters, but Hymenoptera only 58 cm., and Diptera 68 cm.; while
the firefly (Lampyris) can see tolerably well the form of large
objects at a distance of over two meters.

Until further experiments are made, it seems probable, then, that
few if any insects have acute sight, that they see objects best when
moving, and on the whole—except dragon-flies and other predaceous,
swiftly flying insects, such as certain flies, wasps, and bees,
which have very large rounded eyes—insects are guided mainly rather
by the sense of smell than of sight.

=Relation of sight to the color of eyes.=—It appears from the
observations of Girschner that those Diptera with eyes of a uniform
color see better than those with brightly banded or spotted eyes.
Thus those flies (Asilidæ, Empidæ, Leptidæ, Dolichopidæ) whose
predaceous habits requires good or quick sight have uniformly dark
eyes, as have also such flies as live constantly on the wing,
_i.e._, the holoptic Bombyliidæ, Syrphidæ, Pipunculidæ, etc., whose
eyes are also very large.

Those flies whose larvæ are parasitic on other animals have eyes of
a uniform color that they may readily detect the most suitable host
for their young; such are the Bombyliidæ, Conopidæ, Pipunculidæ, and
Tachinidæ.

Certain flies which live in the clear sunlight, as many Dolichopidæ,
some Bombyliidæ, and certain Tabanidæ (Tabanus, Chrysops,
Hæmatopota), and which are often easily caught with the hand, have
eyes spotted or banded with bright or metallic colors. This is also
a sexual trait, as the males of some horse-flies visiting flowers
have eyes of a single color, the spots and bands surviving only on
the lower and hinder parts of the eye, while their voracious
blood-sucking females have the entire eye spotted or banded (Kolbe).

=The color-sense of insects.=—Insects, as Spengel first suggested,
appear to be able to distinguish the color of objects. Lubbock has
experimentally proved that bees, wasps, and ants have this power,
blue being the favorite color of the honey-bee, and violet of ants,
which are sensitive to ultra-violet rays.

It is well known that butterflies will descend from a position high
in the air, mistaking white bits of paper for white flowers; while,
as we have observed, white butterflies (Pieris) prefer white
flowers, and yellow butterflies (Colias) appear to alight on yellow
flowers in preference to white ones.

The late Mr. S. L. Elliott once informed us that on a red barn with
white trimmings he observed that white moths (Spilosoma, Hyphantria,
and _Acronycta oblinita_) rested on the white parts, while on the
darker, reddish portions sat Catocalæ and other dark or reddish
moths. Gross observed that house-flies would frequent a bluish green
ring on the ceiling of his chamber; but if it were covered by white
paper, the flies would leave the spot, though they would return as
soon as the paper ring was removed (Kolbe). We have observed that
house-flies prefer green paper to the yellowish wall of a kitchen,
but were not attracted to sheets of a Prussian blue paper, attached
to the same wall and ceiling.

It is generally supposed that the shape and high colors of flowers
attract insects; but Plateau has made a number of ingenious
experiments which tend to disprove this view. He used in his
investigations the dahlia, with its central head of flowerets, which
contrast so strongly with the corolla. He finds (1) that insects
frequent flowers which have not undergone any mutilation, but whose
form and colors are hidden by green leaves. (2) Neither the shape
nor lively colors of the central head (capitulum) seem to attract
them. (3) The gayly colored peripheral flowerets of simple dahlias
and, consequently, of the heads of other composite flowers, do not
play the rôle of signals, such as has been attributed to them. (4)
The insects are evidently guided by another sense than that of
sight, and this sense is probably that of smell.

LITERATURE ON THE EYES AND VISION

_a_. General

=Serres, Marcel de.= Mémoires sur les yeux composés et les yeux lisses
des insectes. Montpellier, 1813.

=Müller, Johannes.= Zur vergleichenden Physiologie des Gesichtssinnes
der Menschen und der Tiere. 8 Taf. Leipzig, 1826.

—— Ueber die Augen des Maikäfers. (Meckel’s Archiv f. Anat. u. Phys.,
1829, pp. 177–181; Ann. d. Sc. nat., 1829, sér. 1, xviii, pp.
108–112.)

=Dujardin, F.= Sur les yeux simples ou stemmates des animaux
articulés. (C. R. Acad. Sci., Paris, 1847, xxv, pp. 711–714.)

=Gottsche, C. M.= Beitrag zur Anatomie und Physiologie des Auges der
Krebse und Fliegen. (Müller’s Archiv für Anat. u. Phys., 1852, pp.
483–492. Figs.)

=Murray, Andrew.= On insect vision and blind insects. (Edinburgh New
Phil. Jour., new ser. vi, 1857, pp. 120–138.)

=Claparède, Édouard.= Zur Morphologie der zusammengesetzten Augen bei
den Arthropoden. (Zeitschr. f. wissensch. Zool., 1859, x, pp.
191–214, 3 Taf.)

=Dor, H.= De la vision chez les Arthropodes. (Archives Sci. Phys, et
Nat., 1861, xii, p. 22, 1 Pl.)

=Landois, H.= Die Raupenaugen (Ocelli compositi mihi). (Zeitschr. f.
wissensch. Zool., xvi, 1866, pp. 27–44, 1 Taf.)

—— und =W. Thelen.= Zur Entwicklungsgeschichte der fasettierten Augen
von _Tenebrio molitor L._ (Zeitschr. f. wissensch. Zool., xvii,
1867, pp. 34–43, 1 Taf.)

=Schultze, Max.= Untersuchungen über die zusammengesetzten Augen der
Krebsen und Insecten. Bonn, 1868.

=Schmidt, Oscar.= Die Form der Krystallkegel in Arthropodenauge.
(Zeitschr. f. wissensch. Zool., xxx, Suppl., 1878, pp. 1–12, 1 Taf.)

=Grenacher, H.= Untersuchungen ueber das Sehorgan der Arthropoden,
insbesondere Spinnen, Insecten und Crustaceen. (Göttingen, 1879, 4º,
pp. 1–188, 11 Taf.)

=Reichenbach, H.= Wie die Insekten sehen. Fig. (Daheim, xvi Jahrg.,
1880, pp. 284–286.)

=Poletajew, N.= Ueber die Ozellen und ihr Sehvermögen bei den
Phryganiden. (Horæ Soc. Ent. Ross., 1884, xviii, p. 23, 1 Taf. In
Russian.)

=Hickson, S. J.= The eye and optic tract of insects. (Quart. Journ.
Micr. Sc., ser. 2, xxv, 1885, pp. 215–221, 3 Pls.)

=Notthaft, Jul.= Ueber die Gesichtswahrnehmungen vermittelst des
Fazettenauges. (Abhandl. Senckenberg. naturf. Ges., xii., 1880, pp.
35–124, 5 Taf.)

—— Die physiologische Bedeutung des fazettierten Insektenauges.
(Kosmos, 1886, xviii, pp. 442–450, Fig.)

=Mark, E. L.= Simple eyes in arthropods. (Bull. Mus. Comp. Zool.,
1887, xiii, pp. 49–105, 5 Pls.)

=Girschner, E.= Einiges über die Färbung der Dipterenaugen. (Berlin.
Ent. Zeitschr., 1888, xxxi, pp. 155–162, 1 Taf.)

=Graber, V.= Das unicorneale Tracheatenauge. (Archiv f. Mikroskop.
Anat., xvii, 1879, pp. 58–93, 3 Taf.; Nachtrag, p. 94.)

—— Fundamentalversuche über die Helligkeits- und Farbenempfindlichkeit
augenloser und geblendeter Tiere. (Sitzgs.-Ber. Akad. Wissensch.,
Wien, 1883, lxxxvii, pp. 201–236.)

=Dahl, Fr.= Die Insekten können Formen unterscheiden. (Zool. Anz.,
xii, 1889, pp. 243–247.)

=Ciaccio, G. V.= Figure dichiarative della minuta fabbrica degli occhi
de’ Ditteri. Bologna, 1884, 12 Taf., 30 pp.

—— Della minuta fabbrica degli occhi de’ Ditteri. (Mem. Accad.
Bologna, 1886, ser. 4, vi, pp. 605–660.)

—— Sur la forme et la structure des facettes de la cornée et sur les
milieux refringents des yeux composés des Muscidés. (Journ. Micr.,
Paris, 1889, xiii Année, pp. 80–84.)

=Carrière, J.= On the eyes of some invertebrata. (Quart. Journ. Micr.
Sc. 1884, ser. 2, xxiv, pp. 673–681, 1 Pl.)

—— Ueber die Arbeiten von Viallanes, Ciaccio und Hickson. (Biolog.
Centralblatt, v, 1885, pp. 589–597.)

—— Die Sehorgane der Tiere vergleichend anatomisch dargestellt.
München u. Leipzig, 1885, 205 pp., 147 Figs., 1 Taf.

—— Kurze Mitteilungen aus fortgesetzten Untersuchungen über die
Sehorgane. (Zool. Anz., ix Jarhg., 1886, pp. 141–147, 479–481,
496–500.)

=Forel, A.= Les fourmis de la Suisse. (Neue Denkschriften der schweiz.
naturforsch. Gesellsch. xxvi. 1874, pp. 480, 2 Pls.) Separate. pp.
iv u. 457. Genève.

—— Beitrag zur Kenntnis der Sinnesempfindungen der Insekten. (Mitteil.
d. Münchener Ent. Vereins, ii Jahrg., 1878, pp. 1–21.)

—— Sensations des insectes. (Recueil Zool. Suisse, iv, 1886 et 1887.)

=Plateau, F.= L’instinct chez les insectes mis en défaut par les
fleurs artificielles? (Assoc. française avancement des sciences.
Congrès de Clermont. Ferrand, 1876.)

=Plateau, F.= Recherches expérimentales sur la vision chez les
insectes. Les insectes distinguent-ils la forme des objets? (Bull.
Acad. Belg. 3 Sér. x, 1885, pp. 231–250.)

—— Recherches expérimentales sur la vision chez les insectes.

1. Part, _a_. Résumé des travaux effectués jusqu’en 1887 sur la
structure et le fonctionnement des yeux simples. _b_. Vision chez
les Myriapodes. (Ibid. Sér. 3, xiv, 1887, pp. 407–448, 1 Pl.)

3. Part, _a_. Vision chez les chenilles, _b_. Rôle des ocelles
frontaux chez les insectes parfaits. (Ibid. Sér. 3, xv, 1888, pp.
28–91.)

4. Part. Vision à l’aide des yeux composés. _a_. Résumé
anatomo-physiologique. _b_. Expériences comparatives sur les
insectes et sur les vertébrés. (Mém. cour. et autres Mém. Acad.
Belg. 1888, xliii, pp. 1–91, 2 Pls.)

5. Part, _a_. Perception des mouvements chez les insectes. _b_.
Addition aux recherches sur le vol des insectes avenglés. _c_.
Résumé général. (Bull. Acad. Belg. 1888, sér. 3, xvi, pp. 395–457, 1
Pl.)

—— Recherches expérimentales sur la vision chez les Arthropodes, 2 Pls.
(Mém. couronn. et autres Mém. publ. p. l’Acad. Roy. d. Sciences, etc.,
de Belgique, xliii, Bruxelles, 1889.)

=Watase, S.= On the morphology of the compound eyes in the Arthropoda.
(Studies from biol. laborat. Johns-Hopkins Univ., 1890, pp. 287–334, 4
Pls.)

=Stefanowska, M.= La disposition histologique du pigment dans les yeux
des Arthropodes. (Recueil Zool. Suisse, 1890, pp. 151–200, 2 Pls.)

=Pankrath, O.= Das Auge der Raupen und Phryganiden larven. (Zeitschr. f.
wissensch. Zool., 1890, xlix, pp. 690–708, 2 Taf.)

=Lowne, B. Th.= On the modifications of the simple and compound eyes of
insects. (Philos. Trans. Roy. Soc., London, clxix, 1878, pp. 577–602, 3
Pls.)

—— On the structure and functions of the eyes of Arthropoda. (Proc. Roy.
Soc., London, 1883, xxxv, pp. 140–145.)

—— On the compound vision and the morphology of the eye in insects.
(Trans. Linn. Soc., London, 1884, ii, pp. 389–420, 4 Pls.)

—— On the structure of the retina of the blow-fly (_Calliphora
erythrocephala_). (Jour. Linn. Soc., London, 1890, xx, pp. 406–417, 1
Pl.)

=Patten, W.= Eyes of molluscs and arthropods. (Journal of Morphol.,
Boston, 1887, i, pp. 67–92, 1 Pl.; Mitteil. Zool. Stat. Neapel, vi,
1886, pp. 542–756, 5 Taf.)

—— Studies on the eyes of arthropods.—1. Development of the eyes of
Vespa, with observations on the ocelli of some insects. (Ibid., pp.
193–226, 1 Pl.)—2. Eyes of Acilius. (Ibid., 1888, ii., pp. 190–97, 7
Pls.)

—— On the eyes of molluscs and arthropods. (Zool. Anzeiger, 1887, x
Jahrg., pp. 256–261.)

—— Is the ommatidium a hair-bearing sense-bud? (Anatom. Anzeiger, 1890,
v, pp. 353–359, 4 Figs.)

=Exner, S.= Ueber das Sehen von Bewegungen und die Theorie des
zusammengesetzten Auges. (Sitzgsber. d. math. naturwiss. Cl. kais. Akad.
d. Wissens. Wien, lxxii Jahrg., 1875, 3 Abt. Physiologie, pp. 156–190, 1
Taf.)

—— Die Frage von der Funktionsweise der Fazettenauges. (Biolog.
Centralblatt, i, 1881, pp. 272–281.)

—— Das Netzhautbild des Insektenauges. (Sitzgsber. kais. Akad. d.
Wissensch. Wien, 1889, xcviii, 3 Abt., pp. 13–65, 2 Taf. u. 7 Figs.)

—— Durch Licht bedingte Verschiebungen des Pigmentes im Insektenauge und
deren physiologische Bedeutung. (Ibid., pp. 143–151, 1 Taf.)

=Exner, S.= Die Physiologie der fazettierten Augen von Krebsen und
Insekten, 7 Taf., 1, Lichtdruck u. 23 Holzschn. pp. 206. Wien, F.
Deuticke, 1891.

=Lubbock, John.= On the senses, instincts, and intelligence of animals,
with special reference to insects. London, 1888, pp. 292.

=Mallock, A.= Insect sight and the defining power of composite eyes.
(Proc. Roy. Soc., London, 1894, lv, pp. 85–90, 3 Figs.)

_b_. The color-sense

=Nussli, J.= Ueber den Farbensinn der Bienen. (Schweiz. Bienenzeitung,
N. F., ii Jahrg., 1879, pp. 238–240.)

=Kramer.= Der Farbensinn der Bienen. (Ibid., iii Jahrg., 1880, pp.
179–198.)

=Gross, Wilhelm.= Ueber den Farbensinn der Tiere, insbesondere der
Insekten. (Isis v. Russ., v Jahrg., 1880, pp. 292–294, 300–302,
308–309.)

=Lubbock, John.= Ants, bees, and wasps. London, 1882, pp. 448. Also On
the senses, etc., of animals, 1889.

=Graber, Vitus.= Grundlinien zur Erforschung des Helligkeits und
Farbensinnes der Tiere. Prag u. Leipzig, 1884, pp. 322. (See also p.
262.)

=Forel, Auguste.= Les Fourmis perçoisent-elles l’ultra-violet avec
leurs yeux ou avec leur peau? (Arch. Sci. Phys. Nat. Genève, 1886, 3
sér., xvi, pp. 346–350.)

Also the works of Darwin, Wallace, F. Müller, Grant Allen’s The Color
Sense (1879), Beddard’s Animal Coloration, etc.

_b_. The organs of smell

The seat of the organs of smell is mainly in the antennæ, and they may
be regarded as the principal olfactory organs. For our present knowledge
of the anatomy and physiology of the olfactory organs of insects we are
mainly indebted to the recent investigations of Hauser and of Kraepelin.
The following historical and critical remarks are translated from
Kraepelin’s able treatise:

=Historical sketch of our knowledge of the organs of smell.=—In the
first half of the last century began the inquiries as to the seat of
the sense of smell in the arthropods. Thus Réaumur, in his Mémoires
(i, p. 283; ii, 224), expressed the view that in the antennæ was
situated a special organ which might be an organ of smell.

Lesser, Roesel, Lyonet, Bonnet, and others expressed the same
opinion. Before this Sulzer suggested that an “unknown sense” might
exist in the antennæ; others regarded the stigmata as organs of
smell, as these were considered the natural passages for the
olfactory currents. Duméril, in two special treatises as well as in
his Considérations générales, sought to prove the theory as to the
seat of the organs of smell in the stigmata.

Against both of these leading views as to the seat of the sense of
smell were expressed, in the last century, different opinions. Thus
Comparetti thought that the sense of smell might be localized in
very different points of the head, in the antennal club of
lamellicorns, in the sucking-tube of Lepidoptera, in special frontal
holes of flies and Orthoptera, etc., while Bonsdorf considered the
palpi as organs of smell.

Thus four different views, confused, were held at the opening of
this century; the Hamburg zoölogist, M. C. S. Lehrman, in three
different treatises, brought together all the hitherto known
observations and arguments, treated them critically, and completed
them by his own extended studies. Lehrman adopted the opinions of
Reimarus, Baster, Duméril, and Schelver, that the stigmata presented
the most convenient place for the site of the organs of smell.
Cuvier followed throughout the lead of Lehrman, but Latreille
returned to the view of the perception of smell by the antennæ,
while Treviranus considered the mouth of arthropods as the probable
site of the sense of smell, an opinion which, before his time,
Huber, in his experiments on bees, had thought to be correct. Marcel
de Serres (1811) returned again to the palpi, and asserted—at least
in the Orthoptera—their functions to be olfactory, while Blainville,
ten years later, again expressed anew the old opinion that the
antennæ, or at least their terminations, were organs of smell. Up to
that date there was an uncertainty as to the seat of the organs both
of smell and hearing. Fabricius, indeed, had already, in 1783,
thought he had found an organ of hearing at the base of the outer
antenna. In 1826 J. Müller mentioned an already well-known organ in
the abdomen of crickets as an organ of hearing. Müller, however, was
doubtful, from the fact that the nerve passing to this organ arose,
not from the brain, but from the third thoracic ganglion; but,
notwithstanding, he remarks: “Perhaps we have not found the organ of
hearing in insects because we sought for it in the head.” This
discovery was afterwards considerably broadened and extended by
Siebold’s work, for the views of these naturalists on the seat of
both organs had a definite influence, especially in Germany. For
awhile, indeed, Müller’s hypothesis stood in complete contradiction,
so that during the following decennial was presented anew the
picture of opposing observations and opinions as to the nature of
the organs of smell. While Robineau-Desvoidy, at the end of the
twentieth year, and also later, in different writings, strove
energetically for the olfactory nature of the antennæ,
Straus-Dürckheim held fast to the view that the tracheæ possessed
the function under discussion. At the same period Kirby and Spence,
in their valuable Introduction to Entomology, maintained that “two
white cushions on the under side of the upper lip” in the mouth of
biting insects formed a nose or “rhinarium” peculiar to insects.
This opinion was afterwards adopted by Lacordaire (Introduction à
Entomologie), and also by Oken in his Lehrbuch der Naturphilosophie,
while Burmeister, rejecting all the views previously held, believed
that insects might perhaps smell “with the inner upper surface of
the skin.” Müller’s locust’s ear he regarded as a vocal organ.

Besides these occasional expressions of opinion, the French
literature of the thirtieth and fortieth years of this century
recorded a long series of special works, with weighty experimental
and physiological contents, on this subject. Thus Lefebre, in 1838,
described the experiments which he made on bees, and which seemed to
assign the seat of the sense of smell to the antennæ. Dugès reported
similar researches on the Scolopendræ, and Pierret thought that the
great development of the antennæ in the male Bombycidæ might be
similarly interpreted. Driesch sought to give currency to the views
of Bonsdorf, Lamarck, and Marcel de Serres, that the sense of smell
was localized in the palpi, though Duponchel went back to the old
assertion of æroscepsis of Lehrman, _i.e._ of the air-test through
the antennæ, and Goureau again referred the seat of the sense of
smell to the mouth. In England, Newport at this period put forth a
work in which he considered the antennæ as organs of touch and
hearing, and the palpi as organs of smell—a view which, as regards
the antennæ, was opposed by Newman.

Thus the contention as to the use of the antennæ and the seat of the
organs of smell and hearing fluctuated from one side to the other,
and when in 1844 Küster, by reason of his experiments on numerous
insects, again claimed that “the antennæ are the smelling organs of
insects,” he argued on a scientific basis; yet v. Siebold and
Stannius (1848), in their valuable Lehrbuch der vergleichenden
Anatomie (p. 581), remarked that “organs of smell have not yet with
certainty been discovered in these animals.”

The following decennial was of marked importance in the judgment of
many disputed questions. Almost contemporaneously with Siebold and
Stannius’ Lehrbuch appeared an opportune treatise by Erichson, in
which this naturalist first brought forward certain anatomical data
as to the structure of the antennæ of insects. In a great number of
insects Erichson described on the upper surface of the antennæ
peculiar minute pits, “pori,” which, according to him, were covered
by a thin membrane, and to which he ascribed the perception of
smell. A still more thorough work on this subject was published in
the following year by Burmeister, who recognized in the pits of
lamellicorns many small tubercles and hairs; and about the same time
Slater, as also Pierret and Erichson before him had done, out of the
differences of the antennal development in the males and females in
flesh and plant-eating insects, brought together the proof of the
olfactory function of the antennæ. But the most valuable work of
this period is that of Perris, who, after a review of previous
opinions, by exact observations and experiments, a model of their
kind, sought to discover the seat of the sense of smell. He comes to
the conclusion that the antennæ, and perhaps also the palpi, may
claim this sense, and finds full confirmation of Dufour’s views, and
adopts as new the physiological possibility expressed by Hill and
Bonnet, that the antennæ might be the seat of both senses—those of
smell and hearing.

The beautiful works of Erichson, Burmeister, and Perris could not
remain long unnoticed. In 1857 Hicks published complete researches
on the peculiar nerve-endings which he had found in the antennæ,
also in the halteres of flies and the wings of all the other groups
of insects, and which he judged to be for the perception of smell.
But Erichson’s and Burmeister’s “pori” were by Lespès, in 1858,
explained to be so many auditory vesicles with otoliths. This view
was refuted by Claparède and Claus without their deciding on any
definite sense. Leydig first made a decided step in advance. In
different writings this naturalist had busied himself with the
integumental structures of arthropods, and declared Erichson’s view
as to the olfactory nature of the antennal pits as the truest,
before he, in his careful work on the olfactory and auditory organs
of crabs and insects, had given excellent representations of the
numerous anatomical details which he had selected from his extensive
researches in all groups of arthropods. Besides the pits which were
found to exist in Crustacea, Scolopendræ, beetles, Hymenoptera,
Diptera, Orthoptera, Neuroptera, and Hemiptera, and which had only
thus far been regarded as sense-organs, Leydig first calls attention
to the widely distributed pegs and teeth, also considering them as
sense-organs. “Olfactory teeth,” occurring as pale rods, perforated
at the end, on the surface of the antennæ of Crustacea, Myriopoda,
Hymenoptera, Lepidoptera, Coleoptera, are easily distinguished, and
besides the “olfactory pegs” of the palpi, may be claimed as organs
of smell. The nerve-end apparatus first discovered by Hicks in the
halteres and wings, Leydig thinks should be ranked as organs of
hearing.

There was still some opposition to Leydig’s opinion that in the
insects the sense of smell is localized in the antennæ (teeth and
pits), and here the work of Hensen might be mentioned, which in 1860
had a decided influence upon the conclusion of some inquiries.

Thus Landois denied that the antennæ had the sense of smell, and
declared that the pits in the antennæ of the stag beetle were
auditory organs. So, also, Paasch rejected Leydig’s conclusion,
while he sought to again reinstate the old opinion of Rosenthal as
to the olfactory nature of the frontal cavity of the Diptera. In
spite of the exact observations and interesting anatomical
discoveries of Forel in ants, made in 1874, there appeared the great
work of Wolff on the olfactory organs of bees, in which this
observer, with much skill and acuteness, sought to give a basis for
the hypothesis of Kirby and Spence that the seat of the sense of
smell lay in the soft palatine skin of the labrum within the mouth
(_i.e._ the epipharynx). Joseph, two years later, drew attention to
the stigmata as olfactory organs, referring to the olfactory girdle,
and Forel sought by an occasional criticism of Wolff’s conclusions
to prove experimentally the olfactory function of the antenna; but
Graber, in his widely read book on insects, defended the Wolffian
“nose” in the most determined way, and denied to the antennæ their
so often indicated faculty of smell. In 1879 Berté thought he had
observed in the antenna of the flea a distinct auditory organ, and
Lubbock considered the organs of Forel in the antennæ of ants as a
“microscopic stethoscope.” In 1879 Graber described a new
otocyst-like sense-organ in the antennæ of flies, which was
accompanied by a complete list of all the conceivable forms of
auditory organs in arthropods. In this work Graber described in
Musca and other Diptera closed otocysts with otoliths and auditory
hairs, as Lespès had previously done. But Paul Mayer, in two essays,
refuted this view in a criticism of the opinion of Berté, referring
the “otocysts with otoliths” to the well-known antennal pits into
which tracheæ might pass. Mayer did not decide on the function of
the hairs which extend to the bottom of the pits; while in the most
recent research, that of Hauser, the author again energetically
contended for the olfactory function of the antennæ. Both through
physiological experiments and detailed anatomical investigations
Hauser sought to prove his hypothesis, as Pierrot, Erichson, Slater,
and others had done before him, besides working from an evolutional
point of view. In a purely anatomical aspect, especially prominent
are his discovery of the singularly formed nerve-rods in the pits
and peg-like teeth of the Hymenoptera and their development, as well
as the assertion that numerous hairs in the pits described by
Leydig, Meyer, etc., should be considered as direct terminations of
nervous fibres passing into the pits. In the pits he farther, with
Erichson, notices a serous fluid, which may serve as a medium for
the perception of smells. Among the latest articles on this subject
are those of Künckel and Gazagnaire, which are entirely anatomical,
while the latest treatise of Graber on the organs of hearing in
insects opposes Hicks’s theory of the olfactory function of the
nerve-end apparatus in the halteres, wings, etc., and argues for the
auditory nature of these structures. Finally, according to Voges,
the sense of smell is not localized, but spread over the whole body.

My own observations on different groups of insects agree, in
general, with those of Perris, Forel, and Hauser, without being in a
position to confirm or deny the varying relations of the Hemiptera.
That irritating odorous substances (chloroform, acetic acid) cause
the limbs to move in sympathy with the stimulus, I have seen several
times in Acanthosoma; still it may be a gustatory rather than
olfactory stimulus.

Turning now from speculation and simple observation to exact
anatomical and histological data, the nerve-end apparatus seems to
have a distinct reference to the perception of odors. It comprises a
structure composed of nervous substances which are enclosed in a
chitinous tube, and either only stand in relation to the surrounding
bodies by the perforated point, or pass to the surface as free
nerve-fibrillæ.

In insects there is a remarkable and fundamental difference in the
structures of the parts supposed to be the organs of smell. Erichson
was acquainted only with the “pori” covered by a thin membrane; but
Burmeister, in his careful work on the antennæ of the lamellicorns,
distinguished pits at the bottom of which hairs rise from a cup-like
tubercle, from those which were free from hairs. Leydig afterwards
was the first to regard as olfactory organs the so-called pegs
(_kegel_), a short, thick, hair-like structure distinctly perforated
at the tip, which had already, by Lespès in Cercopis, etc., been
described as a kind of tactile papilla. Other very peculiar
olfactory organs of different form, Forel (Fourmis de la Suisse)
discovered in the antennæ of ants, which Lubbock incorrectly
associated with the nerve-end apparatus found by Hicks in other
insects.

As the final result of his researches Kraepelin states that the great
variety of antennal structures previously described may be referred to a
single common fundamental type of a more or less developed free or
sunken hair-like body which stands in connection by means of a wide
pore-canal with a many-nucleated ganglion-cell. The latter sends only a
relatively slender nerve-fibre (axial cord) through the pore-canal into
the hair; but the same is enclosed by epithelial cells which surround
the pore-canal.

Hauser’s researches on the organs of smell in insects were so carefully
made and conclusive that our readers will, we feel sure, be glad to have
laid before them in detail the facts which prove so satisfactorily that
the antennæ of most insects are olfactory rather than auditory in their
functions.

=Physiological experiments.=—First of all one should observe as exactly
as possible the normal animal in its relation to certain odorous
substances, whose fumes possess no corrosive power or peculiarities
interfering with respiration; then remove the antennæ and try after
several days to ascertain what changes have taken place in the relation
of the animal to the substance. In order to come to no false results it
is often necessary to let the insects operated upon rest one or two
days, for immediately after the operation they are generally so restless
that a careful experiment is impossible.

The extirpation of the antennæ is borne by different insects in
different ways; many bear it very easily, and can live for months after
the operation, while others die in the course of a few days after the
loss of these appendages. The animals seem to be least injured if the
operation is performed at a time when they are hibernating. _Pyrrhocoris
apterus_, and many other insects, afforded a very striking proof of this
relation.

Experiments made by placing the antennæ in liquid paraffine so as to
cover them with a layer of paraffine, thus excluding the air, gave the
same result as if the antennæ had been removed.

The experiments may be divided, according to their object, into three
groups. Experiments of the first kind were made on insects in their
relation to strong-smelling substances, as turpentine, carbolic acid,
etc., before and after extirpation of the antennæ. The second group
embraces experiments on the relation of animals as regards their search
for food; and finally the third group embraces experiments on the
relation of the sexes relative to reproduction before and after the
extirpation of the antennæ.

=Relation of insects to smelling substances before and after the loss of
their antennæ.=—Taking a glass rod dipped in carbolic acid and holding
it within 10 cm. of _Philonthus œneus_, found under stones at the end of
February, it was seen to raise its head, turn it in different
directions, and to make lively movements with its antennæ. But scarcely
had Hauser placed the rod close to it when it started back as if
frightened, made a sudden turn, and rushed, extremely disturbed, in the
opposite direction. When he removed the glass rod, the creature busied
itself for some time with its antennæ, while it drew them, with the aid
of its fore limbs, through its mouth, although they had not come into
direct contact with the carbolic acid. There was the same reaction
against oil of turpentine, and it was still more violent against acetic
acid.

After having many times carefully tested the relations of the normal
animal to the substances mentioned, the antennæ were removed from the
socket-cavity.

On the second day after Hauser experimented with the insects, they
exhibited no reaction either against the carbolic acid, the oil of
turpentine, or even against the acetic acid, although he held the glass
rod which had been dipped into it for one or two minutes before and over
the head. The creatures remained completely quiet and immovable, at the
most slightly moving the palpi. They showed otherwise no change in their
mode of life and their demeanor; they ate with great eagerness flesh
which had been placed before them, or dead insects, and some were as
active as usual as late as May. These beetles had, as proved by the
experiments, lost the sense of smell alone; how far the sense of touch
was lost Hauser could not experimentally decide.

The same results followed experiments with species of the genus Ptinus,
Tenebrio, Ichneumon, Formica, Vespa, Tenthredo, Saturnia, Vanessa, and
Smerinthus; also many species of Diptera and Orthoptera, besides Julus
and Lithobius, while many larvæ reacted in the same manner.

Less satisfactory were the experiments with Carabus, Melolontha, and
Silpha; there is no doubt that the species of these genera, through
the extirpation of their antennæ, become more or less injured as to
the acuteness of their powers of smelling; but they never show
themselves wholly unable to perceive strong-smelling substances.

The allurement of the substance acts for a longer time on those
deprived of their antennæ, then they become restless, then they
wander away from the glass tube held before them; still all their
movements are but slightly energetic, and the entire reaction is
indeterminate and enfeebled.

Experiments with the Hemiptera gave still more unfavorable results;
after the loss of their antennæ they reacted to smells as eagerly as
those did which were uninjured.

=Experiments on the use of the antennæ in seeking for food.=—Under this
head experiments were made with Silpha, Sarcophaga, Calliphora, and
Cynomyia.

Silpha and its larva were treated in the following manner: they were
placed in large boxes whose bottoms were covered with moss, etc.; in a
corner of the box was placed a bottle with a small opening, in which was
placed strong-smelling meat. So long as the beetles were in possession
of their antennæ they invariably after a while discovered the meat
exposed in the bottle, while after the loss of their antennæ they did
not come in contact with it.

In a similar way acted the species of Sarcophaga, Calliphora, and
Cynomyia. Hauser, in experimenting with these, placed a dish with a
large piece of decayed flesh on his writing-table. In a short time
specimens of the flies referred to entered through the open window of
the room. The oftener he drove them away from the meat would they swarm
thickly upon it. Then closing the window and catching all the flies, he
deprived them of their antennæ and again set them free. They flew about
the room, but none settled upon the flesh nor tried to approach it.
Where a fly had alighted on a curtain or other object, the decayed flesh
was placed under it so that the full force of the effluvium should pass
over it, but even then no fly would settle upon it.

=Experiments testing the influence of the antennæ of the males in
seeking the females.=—For this purpose Hauser chose those kinds in which
the male antennæ differ in secondary sexual characters from those of the
female, and in which it is known that they readily couple in
confinement, as _Saturnia pavonia_, _Ocneria dispar_, and _Melolontha
vulgaris_. The two first-named insects did not couple after the
extirpation of their antennæ. Of _Melolontha vulgaris_ twenty pairs were
placed in a moderately sized box. On the next morning twelve pairs of
them were found coupling. Hauser then, after removing the first lot,
placed a new set of thirty pairs in the same box, cut off all the
antennæ of the males and those of a number of females. On the following
morning only four pairs were found coupling, and at the end of three
days five others were observed sexually united.

From these experiments Hauser inferred that those insects deprived
of their antennæ were placed in the most favorable situation, such
as they would not find in freedom; for the space in which the
insects moved about was so limited that the males and females must
of necessity meet. But at the same time the results of the
experiments cannot absolutely be regarded as proving that the males,
after the loss of their antennæ, were then not in condition to find
the females, because in the case of the above-mentioned moths, under
similar conditions, after the extirpation of the antennæ no sexual
union took place. If, however, the experiments made do not all lead
to the results desired, Hauser thinks that the results agree with
those of his histological researches, that in the greater number of
insects the sense of smell has its seat in the antennæ. His results
also agree with those of Perris.

=Structure of the organs of smell in insects.=—The olfactory organs
consist, in insects,—_i.e._, all Orthoptera, Termitidæ, Psocidæ,
Diptera, and Hymenoptera, also in most Lepidoptera, Neuroptera, and
Coleoptera,—

1. Of a thick nerve arising from the brain, which passes into the
antennæ.

2. Of a sensitive apparatus at the end, which consists of staff-like
cells, which are modified hypodermis cells, with which the fibres of the
nerves connect.

3. Of a supporting and accessory apparatus, consisting of pits, or peg-
or tooth-like projections filled with a serous fluid, and which may be
regarded as invaginations and outgrowths of the epidermis.

Hauser adds a remark on the distribution of the pits and teeth in the
larvæ of insects, saying that his observations are incomplete, but that
it appears that in the larvæ the teeth are most generally distributed,
and that they occur not on the antennæ alone, but on the palpi; but in
very many larvæ neither pits nor teeth[47] occurred. In the Myriopoda
teeth-like projections occur on the ends of the antennæ. In Lithobius
they form very small, almost cylindrical, pale organs.

[Illustration:

FIG. 268.—Olfactory organ of Caloptenus.
]

[Illustration:

FIG. 269.—Olfactory pits of the antenna of Stenobothrus. This and Fig.
268 after Hauser.
]

LETTERING FOR FIGS. 268, 269, 273, 275, 276, 278–281.—_a_, _a_,
circular thickening of the skin surrounding the opening of the
olfactory pit; _ax_, thread-like continuation of the nerve-cell;
_b_, vesicle-like bottom of the olfactory pit, through which the
olfactory style passes; _br_ bristle in Fig. 283, stout, and
protecting the olfactory pit; _bs_, bent bristle or seta; _ch_,
chitinous integument of the antennæ; _d_, seen in section; _f_,
invaginated pit; _Fv_, Forel’s flask-shaped organ; _Fvo_, its
opening seen from the surface; _gl_, gland-like mass of cells;
_hyc_, hypodermic cells; _i_, entrance into the canal belonging to
the pit; _m_, olfactory membrane; _m′_, _m″_, _mc_, membrane-forming
cell; _n_, nerve of special sense; _nc_, nucleus of the sense- or
ganglion-cell; _o_, opening into the olfactory pit; _p_, olfactory
pit; _cp_, compound pits; _pw_, wall of the pit; _s_, a large seta;
_sc_, sense- or ganglion-cell; _st_, olfactory or sense-style,
sometimes peg-shaped; _tb_, tactile bristle.

[Illustration:

FIG. 270.—_A_, _b_, sense-organ on the abdominal appendages of a fly
(Chrysopila); _c_, sense-organ on the terminal joint of palpus of
Perla.
]

[Illustration:

FIG. 271.—Longitudinal section of part of cercus of _Acheta
domestica_: _ch_, cuticula; _hyp_, hypodermis; _n._ nerve; _h′^1_,
integumental hairs, not sensory; _h^2_, ordinary hair; _h^3_,
sensory hair; _h^4_, bladder-like hair; _sz_, sense-cell.—After Vom
Rath, from Sharp.
]

In the course of a special description of these sense-organs in the
Orthoptera, Hauser describes at length those of _Œdipoda cœrulescens_
and _Caloptenus italicus_. On one antennal joint of Caloptenus (Fig.
268) was often counted 50 pits; on the anterior joints the number
diminishes to about 30. Hauser thinks that in all Orthoptera whose
antennæ are like those of Caloptenus occur similar pits, as he found
them in Stenobothrus (Fig. 269) as well as in Œdipoda. Gryllotalpa
possesses similar pits,—four to six on each antennal joint, making
between 300 and 400 pits on each antenna.[48] In _Mantis religiosa_ the
pits were not detected, but on each joint, except the eighth basal,
there are about 200 small, hollow, curved teeth with a fine opening in
front.

In the Neuroptera (Chrysopa) there occur on the antennæ, besides
numerous very long tactile bristles, small pale, transparent teeth. No
pits could be detected.

In the Hemiptera (two species of Pyrrhocoris only were examined) only
two kinds of tactile bristles occurred, but Hauser detected no pits,
though Lespès states that they are present.

[Illustration:

FIG. 272.—Longitudinal section of apex of palpus of _Pieris brassicæ_:
_sch_, scales; _ch_, cuticula; _hyp_, hypodermis; _n_, nerve; _sz_,
sense-cells; _sh_, sense-hairs.—After Vom Rath, from Sharp.
]

Of the Diptera, Hauser examined more than 60 species. The pits in the
_Diptera brachycera_ (Muscidæ, etc.) are unexceptionally confined to the
third antennal joint. Their number varies extraordinarily in the
different species. _Helophilus florens_ has on each antennal disk only a
single pit, while _Echinomyia grossa_ possesses 200 of them. In flies of
certain families the pits are compound, and contain 10, 20, and often
100 olfactory hairs, partly arising from the coalescence of several
pits. Such pits are usually divided by lateral walls into several
chambers, whose connection is only indicated by their common outlet.
Simple olfactory pits with a single olfactory style were observed only
in the Tabanidæ, Asilidæ, Bombylidæ, Leptidæ, Dolichopidæ, Stratiomyidæ,
and Tipulidæ. In the last the compound forms do not occur at all, but in
the other families mentioned also occur compound pits, receiving from
two to ten nerve-terminations.

The antennal pits of flies are always sac-like invaginations of the
external chitinous integument, of manifold shapes, opening externally
and never closed by a membrane. The pits differ but slightly in the
different species, and that of _Cyrtoneura stabulans_ (Fig. 273) is
described at length as typical of those of brachycerous flies in
general.

The olfactory pits of the Tipulidæ seem to have a somewhat different
structure, since the external passage is closed. It is circular,
surrounded with a slight chitinous wall, and not covered with bristles.
Such pits in their external appearance are like those of the locust
(Caloptenus) and many Hymenoptera. They are situated usually on the
third antennal joint. _Pachyrhina pratensis_ L. has about 60 of them, as
have _Tipula oleracea_ L. and Ctenophora.

In the Lepidoptera, olfactory pits are much like those of flies. Hauser
describes in detail those of _Vanessa io_. Those of the moths were not
examined, but they can be readily and satisfactorily proved to be the
site of the olfactory sense.

[Illustration:

FIG. 273.—Longitudinal section through the third antennal joint of a
fly (_Cyrtoneura stabulans_), showing the compound pits from above
and in section.—After Hauser.
]

[Illustration:

FIG. 274.—Antenna of Adelops, showing the olfactory organs (_p_) in
the five last joints.
]

Historical researches in respect to the Coleoptera generally gave a very
unfavorable result, contrary to Lespès’s views. That author states that
in the Carabidæ the pits are found on the four first joints, but Hauser
could discover them in none which he examined. Usually only tactile
bristles occur, so also in the Cerambycidæ, Curculionidæ, Chrysomelidæ,
and Cantharidæ. In a blind silphid beetle (_Adelops hirtus_) of Mammoth
Cave we have found well-marked olfactory organs (Fig. 274). Similar
organs occur in the antennæ of the Panorpidæ.

Olfactory pits, however, without doubt occur in Silpha, Necrophorus,
Staphylinus, Philonthus, and Tenebrio. The openings of the pits are
small and surrounded with a small chitinous ring; in Silpha,
Necrophorus, and Tenebrio they cannot easily be distinguished from the
insertion-cavities of the bristles, but in Philonthus and Staphylinus
they are less like them, being distinguished by their somewhat larger
size and their often more oval form. In _Philonthus æneus_ about 100
such small pits occur irregularly on the terminal joints; besides, in
this species on each side of the terminal joint is an apparatus which is
like the compound pit generally occurring in the Diptera.

[Illustration:

FIG. 275.—Olfactory pits of the antenna of _Melolontha
vulgaris_.—After Kraepelin.
]

[Illustration:

FIG. 276.—Antennal pit of _Melolontha vulgaris_, seen in vertical
section.—After Hauser.
]

Very remarkable pits occur in the antennal lamellæ of _Melolontha
vulgaris_ (Fig. 275) and other lamellicorns. On the outer surface of the
first and seventh (in the female the sixth) antennal leaf, as also on
the edges of the other leaves, only arise scattered bristles; on the
inner surface of the first and seventh leaves, as also on both surfaces
of the second to sixth leaves, are close rows of rather shallow
depressions of irregular form, some circular, others regularly
hexagonal. Their number is enormous: in the males 39,000, in the females
about 35,000, occur on each antenna.

[Illustration:

FIG. 277.—Organ of smell of Anophthalmus.—After Hauser. _A_, _a_, _b_,
the same in _A. tenuis_, _B_ in _A. tellkampfii_.
]

[Illustration:

FIG. 278.—Section through antennal joint of _Vespa crabro_, showing
the great number of olfactory pits, olfactory and tactile bristles.
_A_, section through an olfactory pit of _Vespa crabro_.—After
Hauser.
]

The antennal pits and teeth of _Dyticus marginalis_ are morphologically
and physiologically identical with those of bees and wasps. In
_Anophthalmus bilimekii_, Hauser found on the last antennal joints about
60 teeth, which essentially differ in form from those previously
described; they are very pale, transparent, cylindrical, elongated, and
bent elbow-shaped on the first third, so that the last two-thirds run
parallel with the antenna. The length of these remarkable teeth is 0.035
mm., their breadth 0.005 mm. He only found them in Anophthalmus, and in
no other species of Carabidæ; they must resemble the teeth described in
Chrysopa. Our species possesses similar processes (Fig. 277). Similar
teeth occur on the maxillary and labial palpi of beetles. _Dyticus
marginalis_ possesses at the end of each terminal palpal joint a group
of very small teeth, which were also detected in _Anophthalmus
bilimekii_, _Melolontha vulgaris_, etc. In _Carabus violascens_ were
detected on the maxillary palpi large, plainly microscopical, white
disks, which are surrounded with a great number of extremely small
teeth.

Whether the above-described organs on the palpi of beetles should be
considered as olfactory or gustatory in their nature can only be
determined by means of physiological experiments; they probably receive
taste-nerve terminations.

[Illustration:

FIG. 279.—Olfactory pits of the antenna of _Vespa vulgaris_.—After
Kraepelin.
]

The Hymenoptera furnished very good material for histological purposes,
so that Hauser could not only study the terminal apparatus of the
olfactory nerves in the perfect insect, but also in three different
stages of the pupa. These are described at length, as regards the
distribution of the pits and teeth, in _Vespa crabro_; each joint of the
antenna (flagellum) possesses between 1300 and 1400 pits, nearly 60
teeth, and about 70 tactile hairs; on the terminal joint there are more
than 200 teeth, so that each antenna has between 13,000 and 14,000
olfactory pits and about 700 teeth (Kegeln). Fig. 278 represents a
cross-section through the penultimate antennal joint of _Vespa crabro_;
we can see how thick are the series of openings on the surface of the
antennæ, and how regular is the distribution of the teeth.

The distribution of the olfactory pits and olfactory teeth is thus seen
to be very general; the deviations are so insignificant that there is no
reason for the establishment of more than one type.

Antennal pits with a small crevice-like opening occur in genera nearly
allied to Vespa and also in most Ichneumonidæ, Braconidæ, and Cynipidæ.
But the crevice-like openings in these families are considerably longer
and often are of a somewhat twisted shape. In all the species with
translucent antennæ we can recognize the inner mouth of the pit as a
round or nearly round disk situated usually under the middle of the
opening. The antennal pits of _Apis mellifica_, as well as those of
Bombus (Fig. 280) and allied genera, differ from those of the
Ichneumonidæ in being not like crevices, but circular openings.

[Illustration:

FIG. 280.—Olfactory pits of the antenna of Bombus.—After Kraepelin.
]

[Illustration:

FIG. 281.—Olfactory pits of the antenna of Formica: _Fv_, Hicks’
“bottle,” Forel’s flask-shaped organ, _Fvo_, its opening.—After
Kraepelin.
]

[Illustration:

FIG. 282.—Supposed olfactory organs at end of antenna of Campodea:
_A_, _C. staphylinus_. _B_, _C. cookei_, from Mammoth Cave.
]

[Illustration:

FIG. 283.—Vertical section through a single olfactory pit in the
antenna of the horse-fly (_Tabanus bovinus_). For lettering see p.
272.—After Hauser.
]

The distribution of the olfactory peg or tooth-like projections seems to
be much more limited than that of the pits in the Ichneumonidæ. Hauser
could not find any. _Apis mellifica_ possesses on each antennal joint
only about twenty slender pale teeth, scarcely a third as many as in
_Vespa crabro_; on the other hand, Formica, of which genus several
species were examined, seems to have far more teeth than pits; they are
relatively long, pale, transparent, and somewhat clavate; they are not
unlike those of Chrysopa; on the terminal joint only occur the round
openings (_Fvo_), which lead into a bottle-shaped invagination of the
integument (_Fv_) and contain an olfactory style (Fig. 281). In the
Tenthredinidæ only teeth and no pits were to be detected. Sirex has on
the under side of the nine last joints of each antenna a group of from
200 to 300 small teeth, which resemble those of _Vespa crabro_; Lyda has
on the terminal joints about 100 teeth. We may add that supposed organs
of smell occur on the antennæ of Campodea (Fig. 282).

Kraepelin also thus briefly summarizes Hauser’s statements as to the
forms of the different organs of smell.

The manifold nature of the antennal organs has, by Hauser, from
thorough studies of the nerve-elements belonging to them, been not
simplified but rendered more complicated. According to this
naturalist we may distinguish the following forms which the
olfactory organs may assume: 1. “Pale, tooth-like chitinous hairs on
the outer surface of the antennæ, which are perforated at the end;
nothing is known as to the relation of the nerve passing into it
(Chrysopa, Anophthalmus). 2. In pit-like depressions of the antennæ
arise _nerve-rods_ (without a chitinous case) which stand in direct
relation with a ganglion-cell lying under it. These pits are either
_simple_, viz. with only an ‘olfactory rod’ (Tabanus, Fig. 283, and
other Diptera, Vanessa), or _compound_ (Muscidæ, and most other
Diptera, and Philonthus). It is important that these pits are partly
_open_ (in the above-named groups of insects), and partly _closed_
and covered with a thin membrane, under whose concavity the
olfactory rods end (Orthoptera, Melolontha, and other lamellicorns).
3. Short, thick pits sunken slightly into the surface of the
antennæ, and over this a chitinous peg perforated at the end, in
whose base, from the interior, projects a very singular nerve-peg,
which is situated over an olfactory ganglion-cell, and provided with
a slender crown of little rods, and flanked on each side by a
flagellum-cell (Hymenoptera). 4. Round or crevice-like pits covered
over by a perforated chitinous membrane with nerve-rods like those
in 3, but in place of the flagellum-cell with ‘membrane-forming’
cells spread before it. Hauser finally mentions further differences
in the ganglion-cells sent out into the nerve-end apparatus. These
exhibit in Diptera and Melolontha only one nucleus, in Hymenoptera a
single very large one (with many nucleoli) and three small ones, in
Vanessa six, in Orthoptera a very large number of nuclei, etc.”

LITERATURE OF THE ORGANS OF SMELL

=Réaumur, Réné Ant. de.= Mémoires pour servir à l’histoire des
insectes. Paris, 1734–42. (i, 283; ii, 224).

=Lesser, F. C.= Insecto-theologia, 1740, p. 262.

=Roesel, A. J.= Insektenbelustigungen, 1746, ii, p. 51.

=Reimarus, H. S.= Allgemeine Betrachtung ueber die Triebe der Thiere
hauptsächlich ihre Kunsttriebe (Instinct). Hamburg, 1760, p. 355.

=Sulzer, J. H.= Die Kenntzeichen der Insecten. Zürich, 1761.

=Lyonet, P.= Traité anatomique de la chenille, 1762, pp. 42, 96, 195.

=Comparetti, A.= De aure interna comparata. Patavii, 1769.

=Bonnet, C.= Œuvres complètes, 1779–1783, ii, p. 36.

—— Contemplation de la nature, Ch. iii, p. 18.

=Scarpa, Ant.= Anatomicæ disquisitiones de auditu et olfactu. Ticini,
1789.

=Huber, F.= Nouvelles observations sur les abeilles, 1792, ii, p. 475.

=Lehrmann, M. C. G.= De antennis insectorum. Londini, Hamburgi, 1799,
p. 48, Diss. posterior; Hamburg and London, 1800, p. 80 (especial
sense, ærocepsis).

=Latreille, P. A.= Histoire naturelle des crustacés et des insectes,
1806–1809, ii, 50.

=Blainville, M. H. D.= Principes d’anatomie comparée, 1822, i, p. 339.

=Dugès, A. L.= Traité de physiologie comparée, 1838, i, p. 161.

=Newport, G.= On the use of the antennæ of insects. (Trans. Ent. Soc.,
London, ii, 1840, pp. 229–248.)

=Robineau-Desvoidy, A. J. B.= Sur l’usage réel des antennes chez les
insectes. (Ann. Soc. Ent. France, 1842, xi Bull., pp. 23–27.)

=Erichson, W. F.= De fabrica et usu antennarum in insectis. Berlin,
1847, 1 Tab., p. 13.

=Perris, E.= Mémoire sur le siège de l’odorat dans les articulés.
(Ann. sc. nat., Sér. 3, 1850, xiv, pp. 159–178.)

=Dufour, L.= Quelques mots sur l’organe de l’odorat et sur celui de
l’ouie dans les insectes. (Actes d. l. Soc. Linn., Bordeaux, 1850,
xvii, Ann. sc. nat., Sér. 3, Zool., xiv, 1850, pp. 179–184.)

=Leydig, F.= Zum feineren Bau der Arthropoden. (Müller’s Archiv, 1855,
pp. 376–480); Lehrbuch der Histologie, 1857, p. 220; Zur Anatomie
der Insekten (Archiv für Anatomie, 1859, pp. 35–89 and 149–183).

—— Ueber Geruchs- und Gehörorgane der Krebse und Insekten. (Archiv f.
Anat. u. Phys., 1860.)

—— Die Hautsinnesorgane der Arthropoden. (Zool. Anzeiger, 1886, pp.
284–291, 308–314, 265–314.)

=Landois, H.= Das Gehörorgan des Hirschkäfers. (Archiv f. mikrosp.
Anat., 1868, iv, pp. 88–95.)

=Troschel, H.= Ueber das Geruchsorgan der Gliedertiere. (Verhandl. d.
naturhist. Vereins d. preuss. Rheinlande u. Westfal., xxvii Jahrg.,
1870, pp. 160–161.)

=Packard, A. S.= The caudal styles of insects sense-organs, _i.e._,
abdominal antennæ. (Amer. Naturalist, 1870, pp. 620, 621. Also Proc.
Bost. Soc. Nat. Hist., 1868, xi, p. 398.)

=Paasch, A.= Von den Sinnesorganen der Insekten im Allgemeinen, von
Gehörund Geruchsorganen im Besondern. (Archiv für Naturgesch., xxxix
Jahrg., i, 1873, pp. 248–275.)

=Chadima, Jos.= Ueber die von Leydig als Geruchsorgane bezeichneten
Bildungen bei den Arthropoden. (Mitteil. d. naturwiss. Ver. f.
Steiermark, 1873, pp. 36–44.)

=Forel, A.= Les Fourmis de la Suisse. (Neue Denkschr. Allg. Schweiz.
Gesellsch. f. d. ges. Naturw., xxvi, 1874, pp. 118, 144.)

—— Études myrmécologiques en 1884, avec une description des organes
sensoriels des antennes. (Bull. Soc. Vaud. sc. nat., 1885, Sér. 2,
xx, pp. 316–380.)

=Graber, V.= Die Insekten. München, 1877.

—— Neue Versuche über die Functionen der Insektenfühler. (Biol.
Centralb., vii, 1887, pp. 13–19.)

=Trouvelot, L.= The use of the antennæ in insects. (Amer. Naturalist,
xi, 1877, pp. 193–196.)

=Mayer, P.= Sopra certi organi di senso nelle antenne dei Ditteri.
(Atti R. Accad. d. Lincei, Roma, Ser. 3, Mem. Cl. sc. fis., mat. e
natur., iii, 1879, p. 11.)

=Hauser, Gustav.= Physiologische und histiologische Untersuchungen
über das Geruchsorgan der Insekten. (Zeitschr. f. wissens. Zool.,
xxxiv, 1880, pp. 367–403, 3 Taf.)

=Kraepelin, Karl.= Ueber die Geruchsorgane der Gliedertiere
(Oster-Programm der Realschule des Johanneums. Hamburg, 1883, pp.
48, 3 Taf.)

=Schiemenz, Paulus.= Ueber das Herkommen des Futtersaftes und die
Speicheldrüsen der Bienen nebst einem Anhange über das Riechorgan.
(Zeitschr. f. wissens. Zool., xxxviii, pp. 71–135, 3 Taf.)

=Plateau, F.= Expériences sur le rôle des palpes chez les arthropodes
maxillés, I., Palpes des insectes broyeurs. (Bull. Soc. Zool.
France, x, 1885, pp. 67–90.)

=Plateau, F.= Une expérience sur la fonction des antennes chez la
Blatte (_Periplaneta orientalis_). (Comptes rend. Soc. Ent. de
Belgique, 1886, pp. 118–122, 1 Fig.)

=Lubbock, J.= On some points in the anatomy of ants. (Monthly Micr.
Journ., 1887, pp. 121–142.)

=Ruland, Franz.= Beiträge zur Kenntniss der antennalen Sinnesorgane
der Insekten. Diss. Marburg, 1888, p. 31, 1 Taf.

=Wasmann, E.= Die Fühler der Insekten. (Stimmen aus Maria-Laach.
Freiburg i. B., 1891, p. 37.)

=Sergi, G.= Ricerche su alcuni organi di senso nelle antenne delle
formiche. (Riv. Filos. Sci. Milano, 1891, p. 10, 3 Figs.)

=Nagel, Wilibald.= Die niederen Sinne der Insekten, 19 Figs.,
Tübingen, 1892, pp. 67.

With the writings of Baster, Lamarck, Cuvier, Treviranus, Oken,
Lefebure, Duméril, Schelver, Bousdorf, Rosenthal, Burmeister,
Slater, Balbiani, Marcel de Serres, Garnier, Berté, Porter, Sazepin,
Reuter, Pierret, Duponchel, Driesch, Küster, Peckham, Lubbock, A.
Dohrn, Lespès.

_c_. The organs of taste

The gustatory organs of insects are microscopic pits or setae, either
hair-like or resembling short pegs, which form the ends of ganglionated
nerves. They are difficult to distinguish morphologically from certain
olfactory structures, and it is owing to their position at or very near
the mouth that they are supposed to be gustatory in nature.

Meinert was the first (1860) to suggest that organs of taste occurred in
ants. He observed in the maxillæ, and tongue of these insects a series
of canals in the cuticula of these organs connected with ganglion-cells,
and through them with the nerves, and queried whether they were not
organs of taste. Forel afterwards (1874) confirmed these observations.
Wolff in an elaborate work (1875) described a group of minute pits (Fig.
284) at the base of the tongue of the honey bee, which he supposed to
possess the sense of smell, but Forel and also Lubbock attributed to
these sensory pits the function of taste. Ten years afterward Will
showed conclusively, both by anatomical studies and by experiments, that
Diptera and Hymenoptera possess gustatory organs. He, however, denied
that the organs of Wolff were gustatory, and maintained that organs of
smell were confined to the maxillæ, paraglossæ, and tongue. As we shall
see, however, what appear to be with little doubt taste-pits, with hairs
or pegs arising from them, are most numerous on the epipharynx of nearly
all insects, and situated at a point where they necessarily must come in
contact with the food as it enters the mouth and passes down the throat.

[Illustration:

FIG. 284.—Taste-pits on the epipharynx (_C_) of the honey-bee: _B_,
horny ridge; _R_, _R_, taste-pits; _L_, _A_, _A_, muscular fibres;
_S_, _S′_, _a b c d e f_, section of skin of œsophagus.—After Wolff.
]

[Illustration:

FIG. 285.—Tip of the proboscis of the honey bee, × 140: _L_, terminal
button or ladle: _Gs_ taste-hairs; _Sh_, guard-hairs; _Hb_, hooked
hairs.—After Will.
]

Kraepelin (1883) discovered taste-organs on the proboscis of the fly,
and taste-hairs at the end of the tongue of the humble-bee (Fig. 285),
and afterwards Lubbock critically discussed the subject, and concluded
that the organs of taste in insects are situated “either in the mouth
itself, or on the organs immediately surrounding it.”

=Structure of the taste-organs.=—The organs have been best studied by
Will, who, besides describing and figuring the chitinous structures,
such as the pits or cups, hairs and the pegs, showed that they were the
terminations of ganglionated nerves.

Figure 286 represents the taste-cups on the maxilla of a wasp, and Fig.
287 the taste-cone or peg projecting from the cup or pit. The cell out
of which the pit and projecting hair or peg are formed is a modified
hypodermis cell; and the seta is apparently a modification of a tactile
hair, situated at the end of a nerve, which just beneath the chitinous
structures passes into a ganglion-cell, which sends off a nerve-fibre to
the main nerve.

Will detected on the tongue of the yellow ant (_Lasius flavus_) from 20
to 24, and in Atta from 40 to 52, of these structures. The number of
pits on the maxillæ vary much, not always being the same on the two
sides of the same insect. We have observed these taste-cups in the honey
and humble bee, not only at the base of the second maxillæ (Fig. 288,
_g_), but also on the paraglossæ (_pg_).

=Distribution in other orders of insects.=—The writer has detected
these taste-cups in other orders than Diptera and Hymenoptera. They
very generally occur in mandibulate insects on the more exposed
surface of the epipharynx (compare pp. 43–46). We have not observed
them in the Synaptera (Lepisma and Machilis).

In the Dermaptera the taste-cups appear to be undeveloped in the
nymph, while in the adult they are fewer in number than in any other
pterygote order yet investigated.

In a species of Forficula from Cordova, Mexico, the taste-pits are
few in number, there being only about a dozen on each side in all;
most of them being situated on the anterior half, and a few near the
base. The taste-pits are provided each with a short fine seta, as
usual arising from the centre.

[Illustration:

FIG. 286.—Under side of left maxilla of Vespa: _Gm_, taste-cups;
_Shm_, protecting hairs; _Tb_, tactile hairs; _Mt_, base of
maxillary palpus.—After Will.
]

In the order Platyptera (including Perla, Pteronarcys, Psocus,
Termes, Eutermes, and Termopsis) we have been unable to detect any
organs of taste.

[Illustration:

FIG. 287.—Section through a taste-cup: _SK_, supporting cone; _N_,
nerve; _SZ_, sense-cell.—After Will. This and Figs. 284–286 from
Lubbock.
]

[Illustration:

FIG. 288.—Tongue of worker honey-bee: _pg_, paraglossæ; _B_, the
same enlarged, showing the taste-papillæ; _C_, _D_, base of a
labial palpus (_mx.′p_) with the taste-papillæ; _E_, taste-cups on
paraglossæ of Bombus; _F_, group of same on left, _G_, on right
side, at base of labial palpus.
]

In the Odonata, however, they are fairly well developed; in
Calopteryx, about 50 taste-cups were discovered; in a species of
Diplax about 28, there being a group of 14 at the base of the
epipharynx on each side of the median line, while in _Æschna heros_
there are two groups of from 25 to 30 taste-cups, situated as in the
two aforenamed genera.

In the Orthoptera the gustatory cups are numerous, well developed,
and present in all the families except the Phasmidæ, where, however,
they may yet be found to occur.

In a large cockroach (Blabera) from Cuba they are well developed. On
each side of the middle of the epipharynx is a curved row of stiff,
defensive spines, and at the distal end of each row is a sensory
field, containing 20 taste-cups on one side and 23 on the other.
Near the front edge of the clypeal region are two more sensory
fields, situated on each side of the median line, there being 35
taste-cups in each field. The taste-cups in this form are rather
smaller than usual in the order.

In the Acrydiidæ they are more numerous than in the Blattidæ. For
example, in _Camnula pellucida_, near what corresponds to the front
edge of the clypeus are two gustatory fields, each bearing about 35
taste-pits. Just in front, under the clypeo-labral suture, are two
similar fields, each containing from 40 to 42 taste-pits. There are
none in front of these. There are thus about 140–150 sense-cups in
all.

The members of the Locustidæ (Fig. 26) appear to be better provided
with the organs of taste than any other Orthoptera, those of the
katydid numbering from 170 to 180. There are from 50 to 60
taste-cups in the front region; behind the middle a group of 25 on
each side, and over an area corresponding to the base of the labrum
and front edge of the clypeus is a sensory field with about 70
taste-cups on each side. They are true cups or beaker-like papillæ,
some with a fine, others with a short, stout, conical seta.

The gustatory organs in the cave cricket (_Hadenœcus subterraneus_,
Fig. 27), from Mammoth Cave, are highly developed, being rounded
papillæ with the nucleus at the top or end. They are grouped on each
side of the middle near the front edge, there being 25 on each side.
An irregular row of these beaker-like organs extends along each
side; some occur under the base of the labrum, but they are most
numerous in a field corresponding to the front edge of the clypeus,
there being 50 on each side, or 100 in all, where in Ceuthophilus
there are only 9 or 10. It would thus appear as if the sense of
taste were much more acute in the cave-dweller than in the
out-of-doors form.

In the Coleoptera taste-cups and setæ are very generally
distributed, though we were unable to detect them in Dendroctonus or
in _Lucanus dama_. As seen in Fig. 57, we have observed numerous
taste-pegs along the maxilla of _Nemognatha lurida_, but otherwise
taste-organs have only been detected in the epipharynx. They not
only occur in the adult beetles, but we have found them in the larvæ
of cerambycid, scarabæid, and other beetles. In the adults
taste-cups appear to be about as well developed in the carnivorous
forms (Carabidæ) as in the phytophagous or lignivorous groups.

In _Chlænius tomentosus_ there are about half as many of these
organs as in Harpalus, while in Calosoma there are 90 taste-cups, 45
on each side, under the base of the labrum. The cups are
papilliform, being rather high, with a seta arising from each.

In the Cicindelidæ, the epipharynx bears a sensory field quite
different from that of the Carabidæ. There are no normal taste-cups,
except a few situated on two large, round, raised areas which are
guarded in front by a few very long setæ. On the surface of each
area are numerous very long setæ which may, if not tactile, have
some other sense, as they arise from cup-like bases or cells. Those
on the outside are like true taste-cups, with a bristle but little
larger than normal in taste-cups generally. We are disposed to
regard this sensory field as a highly specialized gustatory
apparatus.

In the Dyticidæ the taste-cups are nearly as described in the
Carabidæ.

The Staphylinidæ are not well provided with taste-organs. Under the
clypeus of _Staphylinus violaceus_, on each side near the middle, is
a bare rounded area, in which are situated 4 or 5 papilliform
taste-cups, and at the base behind them is another linear group of
about 7 slenderer, somewhat curved, taste-cups. In the Elateridæ
these organs are scantily developed.

In the Buprestidæ (_Buprestis maculiventris_ alone examined) no true
taste-cups were detected. On the other hand, the Lampyridæ are well
supplied with them. Under the clypeus is situated a sensory field
bearing 26 taste-cups, which are rather smaller than usual. Over the
epipharyngeal surface are scattered a few taste-cups, but they are
small and perhaps not gustatory. Under the clypeus of _Lucidota
punctata_ Lec. is a group of 12 taste-cups, and in the middle region
of the epipharynx, situated in a field extending from near the base
to near the front edge, are about 40 taste-cups, which, however, are
not, as is usual, arranged on each side of the median line, but are
scattered among the hairs of the pilose surface of the epipharynx.
In the Cleridæ the taste-cups are few in number.

In the great family of Scarabæidæ, the presence of gustatory organs
is variable. None occur in _Lucanus dama_, though in the June beetle
(_Lachnosterna fusca_ Fröhl.) they are abundantly developed. The
epipharynx bears on each side outside of a spiny area a group of
about 50 taste-cups, each bearing a long seta, those on the outside
of the area passing into a few high, rather slender, papillæ,
without a seta. On the under side of the clypeus is a median group
of 10 taste-cups of singular form, the cups being large, with broad
bases, which posteriorly bear three spines, of which the median one
is the largest.

Taste-cups occur without any known exception in the longicorn
beetles. In _Leptura canadensis_ they are numerous; in _Euryptera
lateralis_ they are abundant along and near the middle of the
anterior half of the labral region, and in _Cyllene robiniæ_ Forst.
(or _pictus_ Drury) they are more numerous than usual, extending in
an unbroken sensory field from near the front margin of the clypeal
region to near the front edge of the epipharynx. The cups vary much
in size, some being one-half as large as others; and those on the
sides of the sensory field bear short, and a few others rather long,
bristles, showing that the taste-cups are modified tactile bristles.

The Tenebrionidæ are fairly well endowed with taste-cups, their
number in _Eleodes obsoleta_ Say amounting to 30 or 40.

Those of the Meloidæ especially are unusual in size and number.

[Illustration:

FIG. 289.—Epipharynx (_ep_) of Nemognatha: _cl_, clypeus; _gh_,
gathering hairs; _tc_, triangular sensory field dotted with taste
cups; _A_, the field enlarged.
]

In _Nemognatha lurida_ Lec. (Fig. 289) the front edge of the
epipharynx contains about 80 remarkably small taste-cups, arranged
irregularly in a triangular sensory space, and not more than ¼ to ⅙,
as large as those on the maxillæ of the same beetle. Unless the
former structures are gustatory it is difficult to account for their
presence here, and it will be observed that the taste-cups in
Epicauta are unusually abundant. Thus in the middle and near the
front of the epipharynx of the blister-beetle over 100 gustatory
cups were counted. They are conical, papilliform, and truncated at
the end as if open, the edge of the opening being ragged, though
bearing no bristle, except in a few cases. Around the edge of the
sinus, on the under side of the labrum, is a regular marginal row of
large, longer, more distinctly chitinized taste-cups, whose walls
are streaked up and down by chitinous thickenings. In _E. callosa_
Lec. there are about 55 taste-cups under the labrum, besides about
10 cells, which may be gustatory structures, situated on either side
of a median setose ridge which passes back under the clypeal region.

The taste-cups of the leaf-beetles are fairly numerous, judging from
an examination of _Diabrotica vittata_. The surface of the
epipharynx is pilose, but the median region is naked, and on the
anterior half bears from 11 to 12 taste-cups, arranged each side of
the median line in a rude Y. On each side, at the base of the labial
region, are two sensitive fields, each bearing about 25 to 26
taste-cups. More were seen under the clypeus.

In the Neuroptera unmistakable taste-cups are not always present. In
_Sialis infumata_ along the median line of the epipharynx and near
the front are about 20 scattered gustatory pegs, which are minute,
but longer and more acute than usual. In _Chauliodes maculatus_
there are one or two taste-cups under the front edge of the clypeus;
others are scattered along the middle from the base of the labrum to
the front, but are not arranged in definite order. In _Corydalis
cornutus_ no sense-cups, pits, or rods are present. In Chrysopa
there are scattered cups armed with a short acute bristle, which are
possibly gustatory in function. In _Myrmeleon diversum_ also the
presence of sense-pits or of taste-cups is doubtful, though a group
of about 12 pits on each side of the clypeal region of the
epipharynx, and a few situated at the base of the labral region, may
be endowed with the sense of taste. In _Mantispa brunnea_, however,
along the middle of the epipharynx are scattered about 30
unmistakable taste-cups, each bearing a short, fine hair.

In the Mecoptera (_Panorpa debilis?_) taste-cups, giving rise to a
minute hair, occur on the labium in two regions, and also on the
maxillæ situated on the stipes near the base of the palpi, and on
the lacinia and galea. They are also to be found on the maxillæ of
_Boreus californicus_, but were not detected on the labium.

They were first detected by Reuter in various microlepidoptera, and
occur on the “basal spot” of the palpi of many butterflies. In a
Tineid moth (_Coleophora coruscipennella_) we have detected what we
suppose to be a group of four taste-pits on the inner side of the
basal joint of the labial palpi.

=Experimental proof.=—No one, says Lubbock, who has ever watched a bee
or wasp can entertain the slightest doubt as to their possession of the
sense of taste. “Forel mixed morphine and strychnine with some honey,
which he offered to his ants. Their antennæ gave them no warning. The
smell of the honey attracted them, and they began to feed; but the
moment the honey touched their lips they perceived the fraud.”

Will at first fed wasps with sugar, so that they frequently visited it;
afterwards he substituted alum for the sugar. Eagerly flying to it, they
had scarcely touched it when they drew back from the distasteful
substance with the most comical gestures, and cleaned their tongues by
frequently running them in and out, repeatedly stroking them with their
fore feet. He noticed a great repugnance to quinine in nearly all the
insects experimented on. Bees and wasps were observed to have a more
delicate gustatory sense than flies, etc., which are more omnivorous in
their tastes.

LITERATURE ON THE ORGANS OF TASTE

=Meinert, F.= Bidrag til de danske myrers naturhistorie. (Kgl. Dansk
Vidensk. selsk. skrifter. Kjoebenhavn. Raekke 5 Naturvid. og math.
Afd., v, 1861, pp. 273–340.)

=Wolff, O. J. B.= Das Riechorgan der Biene. (Nova acta d. K.
Leop.-Carol. Akad., xxxviii, 1875, pp. 1–251, 8 Taf.)

=Joseph, G.= Zur Morphologie des Geschmacksorganes bei Insekten.
(Amtlicher Bericht der 50. Versammlung deutscher Naturforscher und
Aerzte in München, 1877, pp. 227–228.)

=Künckel et Gazagnaire.= Du siège de la gustation chez les insectes
Diptères. Constitution anatomique et physiologique de l’epipharynx
et l’hypopharynx. (Comptes-rend. Acad. Sc., Paris, 1881, xcv, pp.
347–350.)

—— Recherches sur l’organisation et le développement des Diptères et
en particulaire des Volucelles, i, 1875, ii, 1881, 26 Pls.

=Kraepelin, K.= Zur Kenntniss der Anatomie und Physiologie des Rüssels
von Musca. (Zeitschr. f. wissens. Zool., xxxix, 1883, pp. 683–719, 2
Taf.)

=Kirbach, P.= Ueber die Mundwerkzeuge der Schmetterlinge. (Zool.
Anzeiger, 1883, pp. 553–558, 2 Figs.)

=Will, F.= Das Geschmacksorgan der Insekten. (Zeitschr. f. wissens.
Zool., 1885, xlii, pp. 674–707, 1 Taf.)

=Gazagnaire, J.= Du siège de la gustation chez les insectes
Coléoptères. (Comptes-rend. Acad. Sc., Paris, 1886, cii, pp.
629–632; Ann. Soc. Ent. France, Sér. 6, Bull., pp. 79–80.)

=Forel, A.= Expériences et remarques critiques sur les sensations des
insectes. 2^{me} Part. (Recueil Zool. Suisse, iv, 1887, pp.
161–240.)

=Reuter, Enzio.= Ueber den “Basalfleck” auf den Palpen der
Schmetterlinge. (Zool. Anzeiger, 1888, pp. 500–503.)

—— Ueber die Palpen der Rhopaloceren, etc., 6 Taf. Helsingfors, 1896,
pp. 1–578. (Acta Soc. Sc. Fennicæ, xxii, 1896.)

=Packard, A. S.= On the occurrence of organs, probably of taste, in
the epipharynx of the Mecoptera (Panorpa and Boreus). (Psyche, 1889,
v, pp. 159–164.)

—— Notes on the epipharynx, and the epipharyngeal organs of taste in
mandibulate insects. (Psyche, 1889, v, pp. 193–199, 222–228.)

Also Lubbock’s Senses, etc., of animals, and the writings of Briant,
Breithaupt (titles on p. 85).

_d._ The organs of hearing

Although it has been denied by Forel that insects have the sense of
hearing, yet the majority of writers and experimenters agree that
insects are not deaf. On general grounds if, as we know, many insects
produce sounds, it must follow that they have ears to hear, for there is
every reason to suppose that the sounds thus made are, as in other
animals, either for attracting the sexes, for a means of communication,
or to express the emotions. We will begin by briefly describing the
structures now generally supposed to be auditory in function, and about
which there can be no reasonable doubt, and then consider the more
problematical organs, closing with an account of the extremely various
means of producing sounds and cries.

=The ears or tympanal and chordotonal sense-organs of Orthoptera and
other insects.=—The ears or tympana of locusts (Acrydiidæ) are situated
one on each side, on the basal joint of the abdomen, just behind the
first abdominal spiracle. That this is a true ear was first suggested by
J. Müller, and his opinion was confirmed by Siebold, Leydig, Hensen,
Graber, Schmidt, Lubbock, etc.[49]

[Illustration:

FIG. 290.—Ear of a locust (_Caloptenus italicus_), seen from the inner
side: _T_, tympanum; _TR_, its border; _o_, _u_, two horn-like
processes; _bi_, pear-shaped vesicle; _n_, auditory nerve; _ga_,
terminal ganglion; _st_, stigma; _m_, opening, and _m′_, closing,
muscle of the same; _M_, tensor muscle of the tympanum
membrane.—After Graber.
]

The apparatus consists of a tense membrane, the _tympanum_, surrounded
by a horny ring (Fig. 290). “On the internal surface of this membrane
are two horn-like processes (_o_, _u_), to which is attached an
extremely delicate vesicle (_bi_) filled with a transparent fluid, and
representing a membranous labyrinth. This vesicle is in connection with
an auditory nerve (_n_) which arises from the third thoracic ganglion,
forms a ganglion (_ga_) upon the tympanum, and terminates in the
immediate neighborhood of the labyrinth by a collection of cuneiform,
staff-like bodies, with very finely pointed extremities (primitive
nerve-fibres?), which are surrounded by loosely aggregated ganglionic
globules” (Siebold’s Anatomy of the Invertebrates).

[Illustration:

FIG. 291.—Fore tibia of _Locusta viridissima_. _td_, cover of the
drum; _tr_, fissure between the drum and its cover.—After Graber,
from Lang.
]

In the green grasshoppers, katydids, and their allies, the ears are
situated on the fore tibiæ, where these organs can be found after a
careful search (Figs. 291, 292).

The presence of the structure is indicated by the oval disc, the drum,
which is a thin tense membrane covering the auditory apparatus of
nerves, ganglion cells, and auditory rods beneath.

The tympana, or drums, are not present in all Locustidæ and
Gryllidæ, and, as Lubbock states, it is an additional reason for
regarding them as auditory organs, that in those species which
possess no stridulating organs the tympana are also wanting. In many
of the Locustidæ the tympana are covered or protected by a fold of
the skin projecting over them. These covered ones are, Graber
thinks, derived from the open ones.

[Illustration:

FIG. 292.—_A_, fore tibia of a European grasshopper (Meconema),
containing the ear: _Ty_, tympanum or outer membrane; _Tr_ 1, _Tr_
2, tracheæ. _B_, diagrammatic cross-section through the tibia and
ear of the same; _Ty_, tympanum; _Ct_, cuticula; _CM_, hypodermis:
_A_, the auditory organ connecting with the tympanum; _B_,
supra-tympanal auditory organ; _GZ_, the ganglion-cell belonging
to them; _Hst_, the auditory rod connecting with the
ganglion-cells.—After Graber, from Judeich and Nitsche.
]

On examining the apparatus within the leg under the drum, it is seen to
consist of the trachea, the auditory vesicles and rods, ganglion cells,
and acoustic nerve. The trachea is greatly modified (Fig. 292, _Tr_ 1).
On passing into the tibia the trachea enlarges and divides into two
branches, which reunite lower down. The spiracles supplying the air to
this enlarged trachea are considerably enlarged, while in the dumb
species it is of the normal size. The enlarged trachea passes close to
the tympanum, which thus has air on both sides of it: the open air on
the outer, the air of the trachea on its inner surface. In fact, as
Lubbock states, “the trachea acts like the Eustachian tube in our own
ear; it maintains an equilibrium of pressure on each side of the
tympanum, and enables it freely to transmit the atmospheric vibrations.”

[Illustration:

FIG. 293.—The auditory apparatus in the tibia of a grasshopper,
showing the tympanal nerve-endings in situ: _EBI_, terminal vesicles
of Siebold’s organ; _SN_, nerve of the organ of Siebold; _Gr_, group
of vesicles of same; _SO_, nerve-endings of the same; _vT_, front
tympanum; _vTr_, front branch of the trachea; _hT_, hinder tympanum;
_hTr_, hinder branch of the trachea; _Sp_, space between the
tracheæ; _go_, supra tympanal ganglion; _rN_, connecting
nerve-fibrils between the ganglion cells and the terminal vesicles;
_R_, upper, _n-S_, lower, root of the transparent covering membrane.
(Other lettering not explained by author.)—After Graber.
]

[Illustration:

FIG. 294.—Auditory rod of _Gryllus viridissimus_: _fd_, auditory rod;
_ko_, terminal piece.—After Graber, from Lubbock.
]

These tracheæ, says Graber, though formed on a similar plan, present
many variations, corresponding to those of the tympana, and showing
that the tympana and the tracheæ stand in intimate connection with
one another. For instance, in those species where the tympana are
equal, the tracheæ are so likewise; in Gryllotalpa, where the front
tympanum only is developed, though both tracheal branches are
present, the front one is much larger than the other; and where
there is no tympanum, the trachea remains comparatively small, and
even in some cases undivided (Lubbock, _ex_ Graber).

The acoustic nerve, which next to the optic is the thickest in the body,
divides soon after entering the tibia into two branches, one almost
immediately forming a ganglion, the supra-tympanal ganglion, the other
passing down to the tympanum, where it expands into an elongated flat
ganglion, the organ of Siebold (Fig. 293), and closely applied to the
anterior tracheæ.

At the upper part of the ganglion is a group terminating below in a
single row of vesicles, the first few of which are approximately equal,
but which subsequently diminish regularly in size. Each of these
vesicles is connected with the nerve by a fibril (Fig. 293, _vN_), and
contains an auditory rod (Fig. 294). They are said by Graber to be
brightly refractive, hollow (thus differing from the retinal rods, which
are solid), and terminate in a separate end-piece (_ko_). The rods were
first discovered by Siebold, and, as Lubbock remarks, may be regarded as
specially characteristic of the acoustic organs of insects.

[Illustration:

FIG. 295.—Chordotonal organ in nymph of a white ant.—After Müller,
from Sharp.
]

[Illustration:

FIG. 296.—Right half of 8th body-segment of _Corethra plumicornis_:
_g_, ganglion of ventral cord; _lm_, longitudinal muscle; _cn_,
chordotonal nerve; _cl_, chordotonal ligament; _cg_, chordotonal
ganglion; _cs_, rod of chordotonal organ; _cst_, terminal cord;
_tb_, tactile setæ; _hn_, out-going fibres of the integumental
nerves.—After Graber, from Lang.
]

As will be seen in Fig. 293, at the upper part of the tibial organ
of Ephippigera there is a group of cells, and below them a single
row of cells gradually diminishing in size from above downwards.
“One cannot but ask oneself,” says Lubbock, “whether the gradually
diminishing size of the cells in the organ of Siebold may not have
reference to the perception of different notes, as is the case with
the series of diminishing arches in the organ of Corti of our own
ears.”

These organs were supposed to be restricted to the Orthoptera, but
in 1877 Lubbock discovered what seems to resemble the supra-tympanal
auditory organ of Orthoptera in the tibia of the yellow ant (_Lasius
flavus_). Graber confirmed Lubbock’s account, and also discovered
these organs in the tibia of a Perlid (_Isopteryx apicalis_), and
Fritz Müller has detected them in the fore tibiæ of the nymph of
_Calotermes rugosus_ (Fig. 295). To these structures Graber gave the
name of chordotonal organs.

He has also detected these organs in all the legs of other insects
(Trichoptera, Pediculidæ), and auditory rods have been discovered in
the antennæ of Dyticus and of Telephorus by Hicks, Leydig, and
Graber. Graber classifies the chordotonal organs into truncal and
membral. In Coleoptera and Trichoptera they may occur on several
joints of the leg; others are more localized,—thus he distinguishes
femoral (Pediculidæ), tibial (Orthoptera, Perlidæ, Formicidæ), and
tarsal organs (Coleoptera).

A type of chordotonal organ, observed in the body-segments of the
larvæ of several insects by Leydig, Weismann, Graber, Grobben, and
Bolles Lee, is to be seen in the transparent larva of Corethra (Fig.
296), where the auditory organ extends to the skin. It contains at
the point _cs_ two or three auditory rods. In the opposite direction
a fine ligament (_cl_) passes from _cg_ to the skin; in this way the
auditory organ is suspended in a certain state of tension, and is
favorably situated to receive even very fine vibrations. A similar
apparatus has been detected in the larva of Ptychoptera.

=Antennal auditory hairs.=—It is not at all improbable that the antennæ
of different insects contain auditory as well as olfactory structures.
Lubbock has suggested that the singular organs which have only been
found in the antennæ of ants and certain bees, and to which he gives the
name of “Hicks’ bottles” (Fig. 281), may act as microscopic
stethoscopes, while Leydig also regards them as chordotonal organs.

That, however, some of the antennal hairs of the mosquito, as first
suggested by Johnson and afterwards proved experimentally by Mayer, are
auditory, seems well established. Fastening a male mosquito down on a
glass slide, Mayer then sounded a series of tuning-forks. With an Ut_{4}
fork of 512 vibrations per second, some of the hairs were seen to
vibrate vigorously, while others remained comparatively at rest. The
lower (Ut_{3}) and higher (Ut_{5}) harmonics of Ut_{4} also caused more
vibration than any intermediate notes. These hairs, then, are specially
tuned so as to respond to vibrations numbering 512 per second. Other
hairs vibrated to other notes, extending through the middle and next
higher octave of the piano.

Mayer then made large wooden models of these hairs, the one
corresponding to the Ut_{3} hair being about a metre in length, and on
counting the number of vibrations they made when they were clamped at
one end and then drawn on one side, he found that it “coincided with the
ratio existing between the numbers of vibrations of the forks to which
covibrated the fibrils,” or hairs. It should be observed that the song
of the female mosquito corresponds nearly to this note, and would
consequently set the hairs in vibration. Mayer observed that the song of
the female vibrates the hairs of one of the antennæ more forcibly than
those of the other. Those auditory hairs are most affected which are at
right angles to the direction from which the sound comes. Hence from the
position of the antennæ and the hairs a sound will be loudest or most
intense if it is directly in front of the head. If, then, the song of
the female affects one antenna more than another, the male turns his
head until the two antennæ are equally affected, and is thus able to fly
straight towards the female. From his experiments Mayer found that the
male can thus guide himself to within 5° of the direction of the female.
Hence he concludes that “these insects must have the faculty of the
perception of the direction of sound more highly developed than in any
other class of animals.” (Also see Child’s work.)

=Special sense-organs in the wings and halteres.=—Organs of a
special sense, which Hicks supposed to be those of smell, were found
by him near or at the base of the wings of Diptera, Coleoptera, and
less perfect ones in Lepidoptera, Neuroptera, and Orthoptera, with a
trace of them in Hemiptera; but these were considered by Leydig to
be auditory organs, since he found the nerves to end in club-shaped
rods, like those of Orthoptera.

Hicks found, as to the halteres and their sense-organs, that the
nerve in the halter is the largest in the insect, except the optic
nerve; and that at the base of the halteres is a number of vesicles
arranged in four groups, to each of which the nerve sends a branch.
Afterwards Bolles Lee discovered that the vesicles, undoubtedly
perforated, contain a minute hair, those of the upper groups being
protected by hoods of chitin. He regarded them as olfactory organs,
while Lubbock seems inclined to consider them as auditory
structures. Graber also regards the vesicles of Hicks as chordotonal
organs.

In his elaborate account of the balancers, Weinland concludes that
the organs of sense of varying structure occurring at the base of
these appendages allow the perception of movements which the
halteres perform and which enable the fly to steer or direct its
course. The halteres can thus cause differences in the direction of
the flight of a fly in the vertical plane. If the balancers act
unequally, there is a change in direction.

_e._ The sounds of insects

Insects have no true voice; but sounds of different intensity, shrill
cries, and other noises are produced mechanically by insects, either
being love-songs to attract the sexes, to give signals, to communicate
intelligence, or perhaps to express the emotions. The loud, shrill cry
of the Cicada, or chirp of the cricket, is evidently a love-call, and
results in the mating of individuals of separate broods more or less
widely scattered, thus preventing too close interbreeding.

The simplest means of making a noise is that of the death-watch
(Anobium), which strikes or taps on the wall with its head or abdomen.
Longicorn beetles make a sharp sound by the friction of the
mesoscutellum against the edge of the prothoracic cavity, the head being
alternately raised and lowered, Burying-beetles (Necrophorus) rub the
abdomen against the hinder edges of the elytra. Weevils make a loud
noise by rapidly rubbing the tips of the abdomen on the ends of the
elytra.

Landois offers the following summary of the kinds of noises produced
by beetles:

1. Tapping sounds (Bostrycinæ, Anobium).

2. Grating sounds (Elateridæ).

3. Friction without special rasping organs (_Euchirus
longimanus_).

4. Rasping sounds produced by friction:

_a._ Rubbing of the pronotum on the mesonotum (Cerambycidæ
except Spondyli and Prionus).

_b._ Friction of the prosternum on the mesosternum (_Omaloplia
brunnea_).

_c._ Elytra with a rasp at the end (Curculionidæ, Dyticidæ,
Pelobius).

_d._ With a coxal rasp (Geotrupes, Ceratophyus). The male of
Ateuchus stridulates to encourage the female in her work, and
from distress when she is removed. (Darwin.)

_e._ Friction of the edge of the elytra against the femur
(_Chiasognathus grantii_).

_f._ Pygidium with two rasps in the middle (Crioceris, Lema,
Copris, Oryctes, Necrophorus, Tenebrionidæ).

_g._ Abdomen with a grating ridge and four grating plates
(Trox).

_h._ Abdomen with two toothed ridges rubbing on a rasp on edge
of wing-cover (Elaphus, Blethisa, Cychrus).

_i._ Rubbing the elytra on a rasp on the hind wings (_Pelobius
hermanni_).

_j._ Friction of the wing against the abdominal segments
(_Melolontha fullo_).

Mutilla makes a rather sharp noise by rubbing one abdominal segment
against another. Ants (Ponera) have a stridulating apparatus, and
other genera numerous (20) ridges between the segments.

Even certain moths and butterflies emit a rasping or crackling
noise. The death’s-head moth and other sphinges cause it by rubbing
the palpi against the base of the proboscis. These and certain
butterflies are provided with parallel ridges forming a rasp on the
“basal spot” of the inner side of the basal joint of each palpus
(Reuter). A South American butterfly (_Ageronia feronia_) can be
heard for several yards as it flies with a crackling sound. Hampson
finds that the cause of the clicking sound is due to a pair of
strong chitinous hooks attached to the thorax, against which play
the spatulate ends of a pair of hooks attached to the fore wings. An
Australian moth (Hecatesia) flies with a whizzing sound; Vanessa is
said to be sonorous.

The males of Orthoptera produce their shrill cries or chirping
noises, 1, by rubbing the thighs against the sides of the body
(Acrydiidæ); 2, by the friction of the base of the fore wings on
each other (Locustidæ); 3, by rubbing the base of the upper on the
base of the hinder or under pair (Gryllidæ), in the two last there
being a shrilling apparatus consisting of a file on the hind wings,
which rubs on a resonant surface on the fore wings. The females are
not invariably dumb, both sexes of the European Ephippigera being
able to faintly stridulate. Corixa also produces shrill chirping
notes. (Carpenter.)

Certain insects also hum, and have what may perhaps be called a voice.
The cockchafer, besides humming with the wings, produces a sound almost
like a voice. In the large trachea, just behind each spiracle, is a
chitinous process, which is thrown into vibrations by the air during
respiration, and thus produces a humming noise. (Lubbock.) Such is also
the case with flies, the mosquito, dragon-flies, and bees. In flies and
dragon-flies the “voice” is caused by the air issuing from the thoracic
spiracles; while in the humble-bee the abdominal spiracles are also
musical. The sound made by the spiracles bears no relation to that
caused by the wings. Landois tells us that the wing-tone of the
honey-bee is A′; its voice, however, is an octave higher, and often goes
to B″ and C″.

The sounds produced by the wings are constant in each species, except
where, as in Bombus, there are individuals of different sizes; in these
the larger ones generally give a higher note. Thus the comparatively
small male of _Bombus terrestris_ hums on A′, while the large female
hums an entire octave higher.

From the note produced the rapidity of the vibrations can be calculated.
For example, the house-fly, which produces the sound of F, vibrates its
wings 21,120 times in a minute, or 335 times in a second; and the bee,
which makes a sound of A′, as many as 26,400 times, or 440 times in a
second. On the contrary, a tired bee hums on E′, and therefore,
according to theory, vibrates its wings only 330 times in a second.
Marey has confirmed these numbers graphically, and found by experiment
that the fly actually makes 330 strokes in a second. (Lubbock.)

A different kind of musical apparatus is that of the cicada, which has
been elaborately described by Graber. The shrill, piercing notes issue
from a pair of organs on the under side of the base of the abdomen of
the male, these acting somewhat as two kettle-drums, the membrane
covering the depressions being rapidly vibrated.

LITERATURE ON THE ORGANS OF HEARING

_a._ The auditory organs

=Siebold, C. Th. E. von.= Ueber das Stimm- und Gehörorgan der
Orthopteren. (Archiv f. Naturgesch., 1844, x, pp. 52–81.)

=Johnston, Christopher.= Auditory apparatus of the culex mosquito.
(Quart. Journ. Micr. Soc., 1855, iii, pp. 97–102, 1 Fig.)

=Hicks, Braxton.= On a new organ in insects. (Journ. Linn. Soc. Zool.,
London, 1857, pp. 130–140, 1 Pl.)

—— Further remarks on the organ found on the bases of the halteres and
wings of insects. (Trans. Linn. Soc., London, 1857, xxii, pp.
141–145, 2 Pls.)

=Hensen, V.= Ueber das Gehörorgan von Locusta. (Zeitschr. f. wissens.
Zool., xvi, 1866, pp. 190–207.)

=Graber, V.= Bemerkungen über die Gehör- und Stimmorgane der
Heuschrecken und Cicaden (Wiener Sitzungsber. Math.-natur-wiss. Cl.,
lxvi, 1 Abt., 1872, pp. 205–213, 2 Figs.)

—— Die tympanalen Sinnesapparate, der Orthopteren. (Denkschr. d. k.
Akad. d. wissens. Wien, xxxvi, 1876, 2 Abt., pp. 1–140, 10 Taf.)

—— Die abdominalen Tympanalorgane der Cicaden und Gryllodeen. (Ibid.,
1870, xxxvi, pp. 273–290, 2 Taf.)

—— Ueber neue, otocystenartige Sinnesorgane der Insekten. (Archiv f.
mikroskop. Anat., 1878, pp. 35–57, 2 Taf.)

—— Die chordotonalen Sinnesorgane und das Gehör der Insekten. (Archiv
f. mikroskop. Anat., 1882, xx, pp. 506–640; 1883, xxi, pp. 65–145,
Taf.)

=Mayer, Alfred Marshall.= Researches in Acoustics No. 5. 3.
Experiments on the supposed auditory apparatus of the culex
mosquito. (Amer. Jour. Sc. and Arts, Ser. 3, viii, 1874, pp. 81–103;
also Amer. Naturalist, viii, pp. 577–592.)

=Schmidt, Oscar.= Die Gehörorgane der Heuschrecken. (Archiv f.
mikroskop. Anat., xi, 1875, pp. 195–215, 3 Taf.)

=Ranke, J.= Beiträge zu der Lehre von den Uebergangssinnesorganen, das
Gehörorgan der Acridier und das Sehorgan der Hirudineen. (Zeitschr.
f. wissens. Zool., xxv, 1875, pp. 143–164, 1 Taf.)

=Lee, A. Bolles.= Les balanciers des Diptères, leurs organes
sensifères et leur histologie. (Recueil Zool. Suisse, ii, 1885, pp.
363–392, 1 Pl.)

—— Bemerkungen über der feineren Bau der Chordotonalorgane. (Archiv f.
mikroskop. Anat., 1883, xxiii, pp. 133–140, 1 Taf.)

—— Les organes chordotonaux des Diptères et la méthode du chlorure
d’or. (Observations critiques.) (Recueil Zool. Suisse, 1884, ii, pp.
685–689, 1 Pl.)

=Weinland, E.= Ueber die Schwinger (Halteren) der Dipteren (Zeitschr.
f. wissens. Zool., 1890, li, pp. 55–166, 5 Taf.)

=Adelung, N. v.= Beiträge zur Kenntnis des tibialen Gehörapparates der
Locustiden. Inaug. Diss., Leipzig, 1892, 2 Taf.

=Child, Ch. M.= Ein bisher wenig beachtetes antennales Sinnesorgan der
Insekten, mit besonderer Berücksichtigung der Culiciden und
Chironomiden. (Zeitschr. f. wissens. Zool., lviii, 1894, pp.
475–528, 2 Taf.; also, Zool. Anzeiger, xvii Jahrg., pp. 35–38, and
in Annals and Mag. Nat. Hist., 1894 (6), xiii, pp. 372–374).

Also the writings of J. Müller, Kirby and Spence, Burmeister, Gilbert
White, Westwood, Guilding, Meinert, Paasch, Leydig, Viallanes,
Minot, Forel, Mayer, Darwin (Descent of Man, i, ch. x.), F. Müller,
Lubbock (Senses of animals), Westring, Köppen, Bates, Vom Rath,
Peckham, Jourdan, Nagel, etc.

_b._ The sounds made by insects

=Scudder, S. H.= Notes on the stridulation of grasshoppers. (Proc.
Bost. Soc. Nat. Hist., xi, 1868, pp. 306–313 and 316.)

—— The songs of the grasshoppers. (Amer. Naturalist, ii, 1868, pp.
113–120, 5 Figs.)

=Riley, C. V.= The song notes of the periodical Cicada. (Proc. Amer.
Assoc. Adv. Science, xxxiv, 1885, pp. 330–332; also in Kansas City
Rev., October, 1885, pp. 173–175.)

=Swinton, A. H.= (Ent. Month. Mag., 1877.) Sound produced in Ageronia
by a modification of the hook and bristle of the wings.

=Hampson, G. F.= On stridulation in certain Lepidoptera, etc. (Proc.
Zool. Soc. London, 1892, ii, pp. 188–193, Fig; also Psyche, vi, p.
491, 1 Fig.)

With the writings of Landois, Lubbock, Graber, Kolbe, Carpenter (Nat.
Science), Bruyant, and others.

THE DIGESTIVE CANAL AND ITS APPENDAGES

[Illustration:

FIG. 297.—Transverse section through an abdominal segment of larva of
_Megalopyge crispata_, showing the relations of the digestive canal
to the other organs: _int_, hind-intestine, with its mucous or
epithelial layer (_ep_), and _ml_ its outer or muscular layer; _ng_,
ventral ganglion; _ht_, heart; _mp_, urinary tubes; _f_, fat-body;
_sc_, thickened portion of the hypodermis (_hy_) containing the
setigenous cells; _m_, muscles; _m′_, a pair of retractor muscles
inserted near the base of the lateral glandular process (_lgp_);
_cut_, cuticula; _l_, legs. Also compare Figs. 142–144 and 234.
]

[Illustration:

FIG. 298.
The alimentary or digestive canal of insects is a more or less
straight tube, which connects the mouth and anus, the latter
invariably
situated in the last segment of the body, under the last
tergite or suranal plate. It lies directly over the ventral nervous
cord and under the dorsal vessel, passing through the middle of the
body (Fig. 297). It is loosely held in place by delicate retractor
muscles (_retractores ventriculi_, found by Lyonet in the larvæ of
Lepidoptera, and occurring in those of Diptera), but is principally
supported by exceedingly numerous branches of the main tracheæ.
FIG. 298.—Internal anatomy of _Melanoplus femur-rubrum_: _at_, antenna
and nerve leading
to it from the “brain” or supraœsophageal ganglion (_sp_); _oc_,
ocelli, anterior and vertical ones,
with ocellar nerves leading to them from the brain; _œ_, œsophagus;
_m_, mouth; _lb_, labium or under
lip; _if_, infraœsophageal ganglion, sending three pairs of nerves to
the mandibles, maxillæ, and
labium respectively (not clearly shown in the engraving); _sm_,
sympathetic or vagus nerve, starting
from a ganglion resting above the œsophagus, and connecting with
another ganglion (_sg_) near the
hinder end of the crop; _sal_, salivary glands (the termination of the
salivary duct not clearly
shown by the engraver); _nv_, nervous cord and ganglia; _ov_, ovary;
_ur_, origin of urinary tubes;
ovt, oviduct; _sb_, sebaceous gland; _bc_, bursa copulatrix; _ovt_,
site of opening of the oviduct (the
left oviduct cut away); 1–10, abdominal segments. The other organs
labelled in full.—Drawn from
his original dissections by Mr. Edward Burgess.
]

[Illustration:

FIG. 299.—Digestive canal of _Anabrus_: _m_, mouth: _œ_, œsophagus;
_sm_, the sympathetic nerve passing along the crop; _t_, tongue;
_fg_, frontal ganglion; _br_, brain, the nervous cord passing
backward from it; _sr_, salivary reservoir; _sg_, salivary gland;
_pv_, proventriculus; _ur_, origin of urinary tubes; _sb_, sebaceous
gland; 1–10, the ten abdominal segments.—Burgess _del._
]

It is in the higher adult insects differentiated into the mouth and
_pharynx_, the _œsophagus_ or gullet, supplementary to which is the
_crop_ (_ingluvies_) or “_sucking stomach_” of Lepidoptera, Diptera, and
Hymenoptera; the _proventriculus_ or gizzard; the _ventriculus_,
“chyle-stomach,” or, more properly, mid-intestine, and the
hind-intestine, which is divided into the _ileum_, or short intestine,
the long intestine, often slender and coiled, with the colon and the
rectum. Morphologically, however, the digestive or enteric canal is
divided into three primary divisions, which are indicated in the embryo
insect; _i.e._, the _fore-intestine_ (_stomodæum_ of the embryo),
_mid-intestine_ or “chyle-stomach,” and _hind-intestine_ or _proctodæum_
(Fig. 300). The three primary regions, with their differentiations, may
be tabulated thus:—

_Fore-intestine_ (Stomodæum). Mouth and pharynx.
Pumping apparatus of Hemiptera,
Lepidoptera, and Diptera.
Œsophagus.
Crop or ingluvies, food reservoir, or
“sucking stomach.”
Proventriculus.

_Mid-intestine_ (Mesenteron). Mid-intestine, “chylific stomach,” or
ventriculus (with cœcal glands).

_Hind-intestine_ (Proctodæum). Ileum, or short intestine (with the
urinary tubes).
Long intestine.
Colon.
Rectum (with rectal glands).
Anus (with anal glands).

[Illustration:

FIG. 300.—The three primary divisions of the alimentary canal of an
embryonic orthopterous insect: _br_, brain; _sbg_, subœsophageal
ganglion; _ng_, nervous cord; _st_, stomodæum; _pr_, proctodæum;
_mv_, malphigian tubes; _mesen_, mid-intestine; _ht_, heart; _md_,
mandibles; _mx_, _mx′_, 1st and 2d maxillæ.—After Ayers, with some
changes.
]

The appendages of the alimentary canal are: (1) the salivary and poison
glands, which arise from the stomodæum in embryonic life; (2) while to
the chylific stomach a single pair of cœcal appendages (Orthoptera and
larval Diptera, _e.g._ Sciara), or many cœca may be appended; (3) the
urinary tubes, also the rectal glands and the paired anal glands. In a
Hemipter (_Pyrrhocoris apterus_) appendages arise from the intestine in
front of the origin of the urinary tubes. In certain insects a single
cœcal appendage (Nepa, Dyticus, Silpha, Necrophorus, and the
Lepidoptera) arises from the proctodæum.

[Illustration:

FIG. 301—Larva of honey-bee: _g_, brain; _bm_, ventral nervous cord;
_œ_, œsophagus; _sd_, spinning-gland; _cd_, mid-intestine or
chyle-stomach; _ed_, hind-intestine, not yet connected with the
mid-intestine; _vm_, urinary tube; _an_, anus; _st_, stigmata.—After
Leuckart, from Lang.
]

In certain larval insects, as those of the Proctotrypidæ (first larval
stage), the higher Hymenoptera (ichneumons, ants, wasps, and bees, Fig.
301), in the Campodea-like larvæ of the Meloidæ and Stylopidæ, the larva
of the ant-lion (Myrmecoleo), and those of _Diptera pupipara_
(Melophagus), the embryonic condition of the separation of the
proctodæum and mid-gut (mesenteron) persists, the stomach ending in a
blind sac; in such cases the intestine, together with the urinary tubes,
is entirely secretory.

The anus is wanting in the larva of the ant-lion, as also in the wasps
(in which there is a rudimentary colon) and in freshly hatched bees,
though it becomes perfectly formed in the fully grown larvæ (Newport,
art. Insecta, p. 967, and H. Müller).

In the larvæ of lamellicorn Coleoptera (_Melolontha vulgaris_) the
digestive tube is nearly as simple as in bees, though there is a large
colon, which at its beginning forms an immense cœcum, and has also one
anal aperture (Newport).

The length and shape of the digestive canal is dependent on the nature
of the food and also on the mode of life, especially the ease or
difficulty with which the food is digested.

Newport, while stating that the length of the alimentary canal in
larvæ is not in general indicatory of the habits of the species,
makes this qualification after describing the digestive canal of
Calandra as compared with that of Calosoma: “The length and
complication of the intestines, therefore, appear to have some
reference to the quality of the food to be digested, since it is
well known that the food of these latter insects (weevils) is of
difficult assimilation, being as it is chiefly the hard ligneous
fibres of vegetable matter; but they cannot be received as always
indicatory of a carnivorous [or] vegetable feeder, since, as above
remarked, the length of the canal is considerable in one entirely
carnivorous larva, while it is much shorter in some herbivorous, and
particularly in pollenivorous larvæ, as in the Melolontha and the
apodal Hymenoptera.”

[Illustration:

FIG. 302.—Digestive canal of a carabid beetle: _b_, œsophagus; _c_,
crop; _d_, proventriculus; _f_, mid-intestine, or “chyle-stomach,”
with its cœeca; _g_, posterior division of the stomach; _i_, the
two pairs of urinary tubes; _h_, intestine; _k_, rectum; _l_, anal
glands.—After Dufour, from Judeich and Nitsche.
]

Newport also contends that the length of the alimentary canal is not
more indicative in the perfect insect of the carnivorous or
phytophagous habits of the species than in the larva. It is nearly
as long (being from two to three times the length of the whole
body), and is more complicated, in the rapacious Carabidæ (Fig. 302)
than in the honey-sipping Lepidoptera, whose food is entirely
liquid. Referring to the digestive canal of Cicindelidæ, which is
scarcely longer than the body, he claims that “we cannot admit that
the length of the digestive organs, and the existence of a gizzard
and gastric vessels, are indicatory of predacity of habits in the
insect, because a similar conformation of parts exists often in
strictly vegetable feeders. The existence and length of these parts
seem rather to refer to the comparative digestibility of the food
than to its animal or vegetable nature.” Newport then refers to the
digestive canal of Forficulidæ (in which the gizzard is present, the
canal, however, passing in an almost direct line through the body,
making but one slight convolution), “a farther proof that the
_length_ of the canal must not be taken as a criterion whereby to
judge of the habits of a species.” He adds this will apply equally
well to the omnivorous Gryllidæ, in which there exists a short
alimentary canal, but a gizzard of more complicated structure than
that of the Dytiscidæ.

In larval insects and others (Synaptera, Orthoptera, etc.), in which the
digestive canal is simplest, it is scarcely longer than the body, and
passes through it as a straight tube.

In the caterpillar, which is a voracious and constant feeder, the
digestive canal is a large straight tube, not clearly differentiated
into fore-stomach, stomach, and intestine; but in the imago, which only
takes a little liquid food, it is slender, delicate, and highly
differentiated. In the larva the mid-gut forms the largest part of the
canal; in the imago, the intestine becomes very long and coiled into
numerous turns; at the same time the food-reservoir (the “sucking
stomach”) develops, and the excretory tubes are longer.

_a._ The digestive canal

[Illustration:

FIG. 303.—Interior view of the bottom of the head of _Danais
archippus_, the top having been cut away, showing, in the middle,
the pharyngeal sac with its five muscles: the frontal (_f.m_),
dorsal pair (_d.m_), and the lateral pair (_l.m_); _cl_, clypeus;
_cor_, cornea; _œ_, œsophagus; _p.m_, one of the large muscles which
move the labial palp.—After Burgess.
]

It will greatly simplify our conception of the anatomy of the digestive
canal if we take into account its mode of origin in the embryo, bearing
in mind the fact that during the gastrula condition the ectoderm is
invaginated at each pole to form the primitive mouth and fore-gut
(stomodæum) and hind-gut (proctodæum). The cells of the ectoderm secrete
a chitinous lining (intima), which forms the continuation of the outer
chitinous crust, and thus the lining of each end of the digestive canal
is cast whenever the insect molts; while the mid-intestine (mesenteron),
arising independently of the rest of the canal much later in embryonic
life from the mesoderm, is not the result of any invagination, being
directly derived from the mesoderm, and is not lined with chitin.

=The mouth, or oral cavity, and pharynx.=—This is the beginning of the
alimentary bounded above by the clypeus, and labrum, with the
epipharynx, and below by the hypopharynx, or tongue, as well as the
labium. Into it pour the secretion of the salivary glands, which passes
out through an opening at the base of the tongue or hypopharynx. On each
side of the mouth are the mandibles and first maxillæ.

=The sucking or pharyngeal pump.=—This organ has been observed by Graber
in flies and Hemiptera, but the fullest account is that by Burgess, who
was the first to discover it in Lepidoptera. In the milk-weed butterfly
(_Danais archippus_) the canal traversing the proboscis opens into a
pharynx enclosed in a muscular sac (Figs. 303, 304, and 310).

The pharyngeal sac, says Burgess, serves as a pumping organ to suck the
liquid food through the proboscis and to force it backwards into the
digestive canal.

[Illustration:

FIG. 304.—Longitudinal section through the head of Danais, showing the
interior of the left half: _mx_, left maxilla, whose canal leads
into the pharynx; _hph_, floor of the latter, showing some of the
taste-papillæ; _oe_, œsophagus; _ep_, epipharyngeal valve; _sd_,
salivary duct; _d.m_, _f.m_, and _cl_, as in Fig. 302.—After
Burgess.
]

Meinert (“Trophi Dipterorum”) has made elaborate dissections of the
mouth and its armature, including the pharynx of several types of
Diptera, with its musculature. He describes the pharynx as the
principal, and in most Diptera, as the only part of the pump (antlia),
and says: “By the muscles of the pump (_musculis antliæ_) the superior
lamina of the pharynx is varied that the space between the two laminæ
may be increased, and the liquid is thus led through the siphon formed
by the mouth-parts into the mouth” (Fig. 81).

=The œsophagus.=—This is a simple tube, largest in those insects feeding
on solid, usually vegetable, food, and smallest in those living on
liquid food. It usually curves upwards and backwards, passing directly
under the brain, and merges into the crop or proventriculus either at
the back part of the head or in the thorax, its length being very
variable. Its inner walls longitudinally are folded and lined with
chitin.

According to Newport, in the œsophagus of the Gryllidæ, of the two
layers of the mucous lining the second is distinctly glandular and
secretory, and in it there are many thousands of very minute granular
glandular bodies, which probably secrete the “molasses” or repellent
fluid often ejected by these and other insects when captured.

=The crop or ingluvies.=—This, when present, is an enlargement of the
end of the œsophagus, and lined internally with a muscular coat. It is
very large in locusts (Fig. 298), Anabrus (Fig. 299), and other
Orthoptera (the Phasmidæ excepted), in the Dermaptera, and most adult
Coleoptera. A crop-like dilatation in front of a spherical gizzard is
also present in the Synaptera (Poduridæ and Lepismidæ), as well as in
the Mallophaga (Nirmidæ).

[Illustration:

FIG. 305.—Digestive canal of Calandra: _H_, pear-shaped œsophagus;
_I_, crop; _K_, gastric cœca _L_, ilium; _MN_, colon; _P_, urinary
tubes.—After Newport.
]

[Illustration:

FIG. 306.—Section of the crop (_H_), gizzard (_I_), and stomach (_K_)
of Athalia.-After Newport.
]

[Illustration:

_Fig. 307._—Upper side of head and digestive canal of Myrmeleon larva:
_a_, crop; _b_, “stomach”; _c_, free ends of two urinary tubes;
_c′_, common origin of other six tubes; _d_, cœcum; _e_, spinneret;
_ff_, muscles for protruding its sheath; _gg_, maxillary
glands.—After Meinert, from Sharp.
]

In the larvæ of weevils (_Calandra sommeri_) there is a crop (Fig. 305),
but not in the larva of Calosoma; also, according to Beauregard, in the
pollen-eating beetles Zonitis, Sitaris, and Malabris it is wanting,
while in Meloe it is highly developed (Kolbe).

The crop forms a lateral dilatation of the end of the œsophagus in the
larvæ of weevils and of saw-flies (_Athalia centifoliæ_, Fig. 306).

=The “sucking stomach” or food-reservoir.=—This is a thin muscular pouch
connected by a slender neck with the end of the œsophagus or the crop,
when the latter is present. There is no such organ in Orthoptera, except
in Gryllotalpa. It is wanting in the Odonata and in the Plectoptera
(Ephemeridæ); in Platyptera (Perlidæ and Termitidæ), in Trichoptera, and
in Mecoptera (Panorpidæ). In most adult Neuroptera (Myrmeleonidæ,
Hemerobiidæ, and Sialidæ), but not in Rhaphidiidæ, the long œsophagus is
dilated posteriorly into a kind of pouch or crop, and besides there is
often a long “food-reservoir” arising on one of its sides, that of
Myrmeleon (Fig. 307) and Hemerobius being on the right side.

[Illustration:

FIG. 308.—Digestive canal of _Sarcophaga carnaria_: _a_, salivary
gland; _b_, œsophagus; _c_, food reservoir; _f-g_, stomach; _h_,
intestine; _i_, urinary tubes; _k_, rectum.—From Judeich and
Nitsche.
]

A true food-reservoir is present in most Diptera (Fig. 308) as well as
in the larvæ of the Muscidæ, but according to Dufour it is wanting in
some Asilidæ and in _Diptera pupipara_, and according to Brauer in the
Œstridæ. The food-reservoir in Diptera is always situated on the left
side of the digestive canal; there is usually a long neck or canal,
while the reservoir is either oval or more usually bilobed, and often
each lobe is itself curiously lobed.

In Lepidoptera (Figs. 309, 310) the so-called “sucking stomach” is, as
Graber has proved, simply a reservoir for the temporary reception of
food; though generally found to contain nothing but air, Newport has
observed that in flies it is filled with food after feeding. He has
found this to be the case in the flesh fly, and in Eristalis he has
found it “partially filled with yellow pollen from the flowers of the
ragwort upon which the insect was captured,” the pollen grains also
occurring in the canal leading to the bag, in the gullet, and in the
stomach itself. Graber has further proved by feeding flies with a
colored sweet fluid that this sac is only a food-receptacle. As he says:
“It can be seen filling itself fuller and fuller with the colored fluid,
the sac gradually distending until it occupies half the hind-body.”

The food-reservoir of the Hymenoptera is a lateral pouch at the end of
the long, slender œsophagus, and has been seen in the bee to be filled
with honey.

[Illustration:

FIG. 309.—Digestive canal of _Sphinx ligustri_: _h_, œsophagus; _i_,
rudiment of the gizzard; _k_, “stomach”; _q_, its pyloric end; _t_,
food reservoir; _p_, urinary tubes; _l_, ilium; _m_, cœcum of colon;
_n_, rectum; _v_, vent.—After Newport.
]

In the mole-cricket the hinder part of the crop is armed within with
hook-like bristles directed backwards so as not to prevent the energetic
pressure of the food backwards into the proventriculus, and to obviate
the possibility of a regurgitation. (Eberli.)

=The fore-stomach or proventriculus.=—This is especially well developed
in the Dermaptera, in the Orthopterous families Locustidæ, Gryllidæ, and
Mantidæ, while in the Thysanura (Lepisma) there is a spherical gizzard
provided with six teeth. It also occurs in many wood-boring insects, and
in most carnivorous insects, notably the Carabidæ, Dyticidæ, Scolytidæ,
in the Mecoptera (scorpion-flies), in the fleas, and in many kinds of
ants, as well as Cynips, Leucospis, and Xyphidria. It is very muscular,
lined within with chitin, which is usually provided with numerous teeth
arising from the folds. These folds begin in the œsophagus or crop, and
suddenly end where the mesenteron (“chylific stomach”) begins. It has
been compared with the gizzard of birds, and is usually called by German
authors the chewing or masticating stomach. (Kaumagen.)

The proventriculus is best developed in the Gryllidæ (_Acrida
viridissima_), where the six folds at the end of the crop close
together to form a valve between the crop and proventriculus. “They
are each armed with five very minute hooked teeth; and, continued
into the gizzard, develop many more in their course through that
organ. These first teeth are arranged around the entrance to the
gizzard, and seem designed to retain the insufficiently comminuted
food and to pass it on to that organ.

[Illustration:

FIG. 310.—Anatomy of _Danais archippus_ after removal of right half
of the body. _Lettering of the head_: _a_, antenna; _ph_, pharynx;
_pl_, labial palpi; _r_, proboscis; _g_, brain; _usg_,
subœsophageal ganglion. _Lettering of the thorax_: I. II. III.
thoracic segments; _b_{1}_, _b_{2}_, _b_{3}_, the coxal joints of
the three pairs of legs; _bm_, muscles of the wings; _ac_ cephalic
aorta with its swelling; _œ_, œsophagus; _bg_, thoracic ganglia of
the ventral cord; _sd_, salivary glands of one side, those of the
other side cut off near their entrance into the common salivary
duct. _Lettering of the abdomen_: 1–9. abdominal segments; _h_,
heart; _sm_, so-called sucking-stomach (food-reservoir); _cm_,
chyle-stomach; _ag_, abdominal ganglia: _ed_, hind intestine with
colon (_c_) and rectum (_r_); _rm_, urinary vessels; _ov_, ovarial
tubes, those of the right side cut off; _ove_, terminal filaments
of the ovaries; _bc_, bursa copulatrix; _obc_, its outer aperture;
_od_, oviduct; _vag_, vagina; _wo_, its outer aperture; _ad_,
glandular appendages of the vagina partly cut away; _vk_,
connective canal between the vagina and bursa copulatrix with
swelling (receptaculum seminis); _an_, anus.—After Burgess, from
Lang.
]

[Illustration:

FIG. 311.—Transverse section of the proventriculus of _Gryllus
cinereus_: _muc_, muscular walls; _r_, horny ridge between the
large teeth (_sp_).—After Minot.
]

[Illustration:

FIG. 312.—Transverse section of the proventriculus of the
cockroach.—After Miall and Denny.
]

[Illustration:

FIG. 313.—Digestive canal of the honey-bee: _A_, horizontal section
of the body; _lp_, labial palpus; _mx_, maxilla; _e_, eye; _pro.
t_, prothorax; _mesa. t_, mesothorax; _meta. t_, metathorax; _dv_,
dorsal vessel; _v_, _v_, ventricles of the same; No. 1, No. 2, No.
3, salivary gland systems; _œ_ œsophagus; _g_, _g_, ganglia of
chief nerve-chain; _n_, nerves; _hs_, honey-sac; _p_, petaloid
stopper or calyx of honey-sac or stomach-mouth; _c. s_, chyle
stomach; _bt_, urinary tubes; _si_, small intestine (ilium); _l_,
“lamellæ or gland-plates of colon,” rectal glands; _li_, rectum.
_B_, cellular layer of stomach; _gc_, gastric cells, × 200. _C_,
urinary tube; _bc_, cells; _t_, trachea. _D_, inner layer, with
gastric teeth (_gt_).—After Cheshire.
]

“Next to these in succession on each of the longitudinal ridges are
four flat, broad, somewhat quadrate teeth, each of which is very
finely denticulated along its free margin. These extend about
half-way through the gizzard. They appear to be alternately elevated
and depressed during the action of the gizzard, and to serve to
carry on the food to the twelve cutting teeth, with which each ridge
is also armed, and which occupy the posterior part of the organ.
These teeth are triangular, sharp-pointed, and directed posteriorly,
and gradually decrease in size in succession from before backward.
Each tooth is very strong, sharp-pointed, and of the color and
consistence of tortoise shell, and is armed on each side by a
smaller pointed tooth. These form the six longitudinal ridges of the
gizzard, between each two of which there are two other rows of very
minute teeth of a triangular form, somewhat resembling the larger
one in structure, occupying the channels between the ridges. The
muscular portion of the gizzard is equally interesting. It is not
merely formed of transverse and longitudinal fibres, but sends from
its inner surface into the cavity of each of the large teeth other
minute but powerful muscles, a pair of which are inserted into each
tooth. The number of teeth in the gizzard amounts to 270, which is
the same number in these Gryllidæ as found formerly by Dr. Kidd in
the mole-cricket. Of the different kinds of teeth there are as
follows: 72 large treble teeth, 24 flat quadrate teeth, 30 small
single-hooked teeth, and 12 rows of small triangular teeth, each row
being formed of 12 teeth. This is the complicated gizzard of the
higher Orthoptera.” (Newport.)

In the more generalized cockroach, there are six principal folds,
the so-called teeth, which project so far inwards as to nearly meet
(Fig. 312). The entire apparatus of muscles and teeth is, as Miall
and Denny state, “an elaborate machine for squeezing and straining
the food, and recalls the gastric mill and pyloric strainer of the
crayfish. The powerful annular muscles approximate the teeth and
folds, closing the passage, while small longitudinal muscles, which
can be traced from the chitinous teeth to the cushions, appear to
retract these last, and open a passage for the food.”

As in the fore-stomach or proventriculus of the lobster, the solid,
rounded teeth do not appear to triturate the solid fragments found in
the organ, but act rather as a pyloric strainer to keep such bodies out
of the chylific stomach. We accept the view of Plateau that this section
of the digestive canal in insects, which he compares to the psalterium
of a ruminant, is a strainer rather than a masticatory stomach, and both
Forel and Emery, as well as Cheshire, take this view.

The proventriculus of the honey-bee (Fig. 313, _hs_) is called by
apiarians the “honey-sac” or “honey-stomach.” Cheshire states that if it
be carefully removed from a freshly killed bee, its calyx-like
“stomach-mouth” may be seen to gape open and shut with a rapid snapping
movement. The entrance to the stomach is guarded by four valves, each of
which is strongly chitinous within, and fringed along its edge with
downward-pointing fine stiff bristles. By the contraction of the
longitudinal muscles (_lm_), the valves open to allow the passage of
food from the honey-sac to the “chyle-stomach.” It is closed at will by
circular muscles (_tm_). Then the bee can carry food for a week’s
necessities, either using it rapidly in the production of wax, or eking
it out if the weather is unfavorable for the gathering of a new store.

[Illustration:

FIG. 314.—“Honey-sac stopper,” “stomach-mouth,” or calyx-bell of
honey-bee, × 50. _A_, front view of one of the lobes of the
calyx-bell; _l_, lip-like point, covered by down-turned bristles
(_b_); _sm_, side membrane. _B_, longitudinal section of the
stomach-mouth, with continuations into entrance of chyle-stomach;
_l_, _l_, lip-like ends of leaflets; _s_, setæ; _lm_, longitudinal
muscles; _tm_, transverse muscles in cross-section; _cl_, cell-layer
of honey-sac; _LM_, _TM_, longitudinal and transverse muscles of
same; _nc_, nucleated cells of tubular extension of stomach-mouth
into chyle-stomach; _lm′_, _tm′_, longitudinal and transverse
muscles of chyle stomach; _c_, _c_, cells covered within by an
intima. _C_, cross-section of stomach-mouth; _m_, cross-section of
muscles seen at _lm_ in _B_; _tm_, transverse muscles surrounding
stomach-mouth. _D_, cross-section through small intestine; _a_ and
_m_, longitudinal and surrounding muscles.—After Cheshire.
]

Cheshire also shows that when bees suck up from composite and other
flowers nectar together with much pollen, the outside wrinkled
membrane (_sm_, _A_, Fig. 314) “is seen to continually run up in
folds, and gather itself over the top of the stomach-mouth, bringing
with it, by the aid of its setæ, the large pollen-grains the nectar
contains.” The lips (_l_, _l_, _B_, Fig. 314), now opening, take in
this pollen, which is driven forwards into the cavity made between
the separating lips by an inflow of the fluid surrounding the
granules. The lips in turn close, but the down-pointing bristles are
thrown outwards from the face of the leaflet, in this way revealing
their special function, as the pollen is prevented from receding
while the nectar passes back into the honey-sac, strained through
between the bristles aforesaid, the last parts escaping by the
loop-like openings seen in the corners of _C_, Fig. 314. The whole
process is immediately and very rapidly repeated, so that the pollen
collects and the honey is cleared. “Three purposes, in addition to
those previously enumerated, are thus subserved by this wondrous
mechanism. First, the bee can either _eat or drink_ from the mixed
diet she carries, gulping down the pollen in pellets, or swallowing
the nectar as her necessities demand. Second, when the collected
pollen is driven forwards into the chyle-stomach, the tube
extension, whose necessity now becomes apparent, prevents the
pellets forming into plug-like masses just below _p_, Fig. 313, for,
by the action of the tube, these pellets are delivered into the
midst of the fluids of the stomach, to be at once broken up and
subjected to the digestive process. And third, while the little
gatherer is flying from flower to flower, her stomach-mouth is busy
in separating pollen from nectar, so that the latter may be less
liable to fermentation and better suited to winter consumption. She,
in fact, carries with her, and at once puts into operation, the most
ancient, and yet the most perfect and beautiful, of all
‘honey-strainers.’”

Forel’s experiments on the proventriculus of ants prove that through
its valvular contrivance it closes the passage from the crop to the
mid-intestine (“chylific stomach”), and allows the contents of the
former to pass slowly and very gradually into the latter. Emery
confirms this view, and concludes that the organ in the Camponotidæ
and in the Dolichoderidæ provided with a calyx-bell, usually
regarded as a triturating stomach (Kaumagen), but more correctly as
a pumping stomach, consists of parts which perform two different
functions. Under the operation of the muscles of the crop the
entrance to the pumping stomach becomes closed, in order by such
spasmodic contraction to prevent the flow of the contents of the
crop into the proventriculus. By the pressure of the transverse
muscles of the proventriculus its contents are emptied into the
mid-intestine, while simultaneously a regurgitation into the crop is
prevented. In the Dolichoderidæ and Plagiolepidinæ the closure in
both cases is effected by the valves. In the true Camponotidæ there
are two separate contrivances for closing; the calyx belonging to
the crop-musculature, while the valves essentially belong to the
proventricular pumping apparatus.

Opinions vary as to the use of this portion of the digestive canal.
Graber compares it to the gizzard of birds, and likens the action of
the rosette of teeth to the finer radiating teeth of the sea-urchin,
and styles it a chopping machine, which works automatically, and
allows no solid bits of food to pass in to injure the delicate walls
of the stomach (mid-gut).

He also states that the food when taken from the proventriculus is
very finely divided, while that found in the œsophagus contains
large bits.

Kolbe says that this view has recently been completely abandoned,
and that the teeth are used to pass the food backwards into the
chylific stomach. “But Goldfuss had denied the triturating action of
the proventriculus of the Orthoptera (Symbolæ ad Orthopterorum
quorundam Œconomiam, 1843), stating that the contents of the same
are already fluid in the gullet, so that the fore-stomach (Kaumagen)
does not need to comminute the food” (Kolbe). In the Gryllidæ and
Locustidæ, just before the posterior opening of the proventriculus
into the stomach the chitinous lining swells into a ring and
projects straight back as the inner wall of the cylindrical chylific
stomach. The muscular layer forms two sac-like outgrowths or folds,
which separate on the circular fold from the chitinous membrane.
This apparatus only allows very finely comminuted food to pass into
the stomach.

In the Acrydiidæ (_Eremobia muricata_) at the end of the
proventriculus, where it passes into the stomach, is a small
circular fold which hangs down like a curtain in the stomach.

=The œsophageal valve.=—Weismann[50] states that the origin of the
proventriculus in the embryo of flies (Muscidæ) shows that it should be
regarded as an intussusception of the œsophagus. While in the embryo the
invaginated portion of the œsophagus is short, after the hatching of the
larva it projects backwards into the mid-intestine. Kowalevsky also
observed in a young muscid larva, 2.2 mm. in length, that the œsophagus,
shaped like a tube, extends back into the expanded portion
(proventriculus) and opens into the stomach (Fig. 315, _A_). In a larva
10 mm. long the funnel is shorter, the end being situated in the
proventriculus (Fig. 315, _B_, _pr_). In the cavity between the outer
(_o_) and inner wall (_i_) no food enters, and the use of this whole
apparatus seems to be to prevent the larger bits of food from passing
into the chylific stomach (Kowalevsky).

[Illustration:

FIG. 315.—Œsophageal valve of young muscid larva: _m_, its opening:
_t_, thickening of the cells; _mes_, mesoderm.—After Kowalevsky.
]

Beauregard has found a similar structure in the Meloidæ, and calls
it the “cardiac valvule” (Fig. 318, _Kl_). It was observed by
Mingazzini in the larvæ of phytophagic lamellicorn beetles, and
Balbiani described it in a myriopod (Cryptops) under the name of the
“œsophageal valvule.”

Gehuchten describes a homologous but more complicated structure in a
tipulid larva (_Ptychoptera contaminata_), but differing in
containing blood-cavities, as a tubular prolongation of the
posterior end of the œsophagus which passes through the
proventriculus and opens at various positions in the anterior part
of the chylific stomach (Fig. 316).

The three layers composing this funnel are distant from each other
and separated by blood-cavities, the whole forming “an immense
blood-cavity extended between the epithelial proventricular lining
and the muscular coat.”

According to Schneider the longitudinal muscular fibres of the fore
and hind gut in insects pass into the stomach (mid-gut). The
anterior part of the fore-gut has generally only circular fibres.
When, however, the longitudinal fibres arise behind the middle, then
they separate from the digestive canal and are inserted a little
behind the beginning of the chylific stomach. Hence there is formed
an invagination of the proventriculus, which projects into the
cavity of the stomach.

Schneider describes this process, which he calls the “beak,” as an
invagination of the fore-stomach which projects into the cavity of
the stomach. The two layers of the invagination in growing together
form a beak varying in shape, being either simple or lobed and armed
with bristles or teeth. This beak is tolerably large in Lepisma,
Dermaptera (Forficula), Orthoptera, and in the larvæ and adults of
Diptera, but smaller in the Neuroptera and Coleoptera, while in
other insects it is wanting.

=Proventricular valvule.=—Gehuchten also describes in Ptychoptera
what he calls “the proventricular valvule,” stating that it is “a
circular fold of the intestinal wall” (Fig. 310, _vpr_). He claims
that it has not before been found, the “proventricular beak” of
Schneider being regarded by him as the œsophageal valvule.

[Illustration:

FIG. 316.—Digestive canal of _Ptychoptera contaminata_: _gs_, salivary
glands; _ra_, œsophagus; _pr_, proventriculus; _gt_, crown of eight
small tubular glands; _im_, mid-intestine; _ga_, two accessory white
glands; _vm_, urinary vessels; _ig_, small intestine; _gi_, large
intestine; _r_, rectum; _A_, the proventriculus in which the hinder
end of the œsophagus extends as far as the chyle-stomach. _B_,
longitudinal section of the proventricular region; _sph_, muscular
ring or œsophageal sphincter; _ppr_, wall of the proventriculus;
_e_, circular constriction dividing the cavity of the proventriculus
in two; _vpr_, circular fold of the wall of the mid-intestine
forming the proventricular valve; _vœ_, œsophageal valve.—After
Gehuchten.
]

=The peritrophic membrane.=—This membrane appears first to have been
noticed by Ramdohr in 1811 in _Hemerobius perla_. It has been found by
Schneider, who calls it the “funnel.” On the hinder end of the
fore-stomach, he says, the cuticula forms a fold enclosing the outlet of
the fore-stomach, and extending back like a tube to the anus. This
“funnel,” he adds, occurs in a great number of insects. It has been
found in Thysanura, but is wanting in Hemiptera. In the Coleoptera it is
absent in Carabidæ and Dyticidæ. It is generally present in Diptera and
in the larvæ of Lepidoptera, but not in the adults. In Hymenoptera it
has been found in ants and wasps, but is absent in Cynipidæ,
Ichneumonidæ, and Tenthredinidæ. All those insects (including their
larvæ) possessing this funnel eat solid, indigestible food, while those
which do not possess it take fluid nourishment. It is elastic, and
firmly encloses the contents of the digestive tract. Until Schneider’s
discovery of its general occurrence, it had only been known to exist in
the viviparous Cecidomyia larvæ (Miastor). Wagner, its discoverer,
noticed in the stomach of this insect a second tube which contained
food. Pagenstecher was inclined to regard the tube as a secretion of the
salivary glands. Metschnikoff, however, more correctly stated that the
tube consisted of chitin, but he regarded it as adapted for the removal
of the secretions. (Schneider.) Plateau, however, as well as Balbiani,
the latter calling it the “peritropic membrane,” considers this membrane
as a secretion of the chylific stomach, and that it is formed at the
surface of the epithelial cells. It surrounds the food along the entire
digestive tract, forming an envelope around the fæcal masses. On the
other hand, Gehuchten states that in the larva of Ptychoptera its mode
of origin differs from that described by Plateau and by Schneider, and
that it is a product of secretion of special cells in the
proventriculus.

=The mid-intestine.=—This section of the digestive canal, often, though
erroneously, called the “chylific stomach” or ventriculus, differs not
only in its embryonic history, but also in its structure and physiology
from the fore and hind intestine of arthropods, and also presents no
analogy to the stomach of the vertebrate animals. In insects it is a
simple tube, not usually lined with chitin, since it is not formed by
the invagination of the ectoderm, as are the fore and hind intestine,
the absence of the chitinous intima promoting the absorption of soluble
food. Into the anterior end either open two or more large cœcal tubes
(Fig. 299), or its whole outer surface is beset with very numerous fine
glandular filaments like villi (Fig. 317 and Fig. 329).

The mid-intestine varies much in size and shape; it is very long in the
lamellicorn beetles (Melolontha and Geotrupes), and while in Meloë it is
very large, occupying the greatest part of the body-cavity, in the
longicorn beetles and in Lepidoptera it is very small. The pyloric end
consists of an internal circular fold projecting into the cavity. In the
Psocidæ (Cæcilius) the pyloric end is prolonged into a slender tube
nearly as long as the larger anterior portion.

The limits between the mid and hind intestine are in some insects
difficult to define, the urinary tubes sometimes appearing to open into
the end of the mid-intestine (“stomach”). The latter also is sometimes
lined with an intima. The limits are also determined by a circular
projection, directly behind which is an enlargement of the intestine in
the shape of a trench (_rigole_), or circular _cul-de-sac_ (the “pyloric
valvule” of some authors, including Beauregard), while the walls of the
small intestine contract so as to produce a considerable constriction of
the cavity of the canal. This constriction exactly coincides with the
beginning of the double layer of circular muscles in the wall of the
small intestine. An internal layer, which is the continuation of the
circular muscles of the chylific stomach, and an external layer much
more developed probably belong to this part of the alimentary canal.
Since the homologue of the circular fold occurs in the locust as well as
in Diptera, it is probably common to insects in general.

[Illustration:

FIG. 317.—Digestive canal of _Carabus monilis_: _h_, œsophagus; _i_,
gizzard or proventriculus; _k_, “stomach,” with its cœca (_r_); _p_,
urinary tubes; _q_, their point of insertion; _m_, _n_, colon, with
cœcal glands; _s_, anal glands; _a_, _b_, _c_, a gastric cœcum; _a_,
_b_, portion of lining of gizzard.—After Newport.
]

[Illustration:

FIG. 318.—Digestive canal of Meloe: _sch_, œsophagus; _Kl_, œsophageal
valve; _mD_, mid-intestine; _eD_, hind-intestine; _Ei_, eggs; _g_,
sexual opening.—After Graber.
]

Gehuchten adds that the limit set by the circular projection does
not exactly coincide with the opening into the intestine of the
urinary tubes and the two annexed glands. He shows by a section (his
Fig. 133) that the tubular glands open into the alimentary canal in
front of the circular fold. It is the same with the Malpighian
tubes. They are not, therefore, he claims, dependences of the
terminal intestine, but of the mid-intestine. Beauregard has
observed the same thing in the vesicating insects (Meloidæ). The
Malpighian tubes, he says, open into the “chylific stomach” before
the valvular crown. This arrangement does not seem to be general,
because, according to Balbiani, the Malpighian vessels open into the
beginning of the intestine in Cryptops. Compare also Minot’s account
of the valve in locusts separating the stomach from the intestine,
and in front of which the urinary or Malpighian tubes open.

=Histology of the mid-intestine.=—The walls of the stomach are composed
of an internal epithelium, a layer of connective tissue, with two
muscular layers, the inner of which is formed of unstriated circular
muscular fibres, and the outer of striated longitudinal muscular fibres.

In the cockroach short processes are given off from the free ends of the
epithelial cells, as in the intestine of many mammals and other animals.
“Between the cells a reticulum is often to be seen, especially where the
cells have burst; it extends between and among all the elements of the
mucous lining, and probably serves, like the very similar structure met
with in mammalian intestines, to absorb and conduct some of the products
of digestion.” (Miall and Denny.)

Gehuchten shows that the epithelial lining of the mesenteron (chylific
stomach) of the dipterous larva Ptychoptera is composed of two kinds of
cells, _i.e._ secreting or glandular cells and absorbent cells, the
former situated at each end of the stomach, and the absorbent cells
occupying the middle region. The part played by these cells in digestion
will be treated of beyond in the section on digestion. (See p. 327.)

=The hind-intestine.=—In many insects this is divided into the ileum, or
short intestine, and the long intestine. The limit between the intestine
and stomach is externally determined by the origin of the urinary tubes,
which are outgrowths of the anterior end of the proctodæum. Like the
fore-intestine the hind-intestine is lined with a thick muscular layer,
and, as Gehuchten states, the passage from the epithelial lining of the
stomach (mid-intestine) to the muscular lining of the intestine is
abrupt.

=Large intestine.=—In Ptychoptera, as described by Gehuchten, there are
no precise limits between the small and large intestine; the epithelium
of the large intestine has a special character, and its constituents
present a close resemblance to the absorbed cells of the chylific
stomach, being like them large and polygonal. The muscular layer is not
continuous, and is formed of longitudinal and circular fibres, the
latter being the larger.

=The ileum=—Though in most insects slender, and therefore called the
small intestine, the ileum is in locusts (Fig. 298) and grasshoppers
(Anabrus, Fig. 299) as thick as the stomach. In many carnivorous beetles
(Dyticus, Fig. 320, _il_, and Necrophorus) it is very long, but rather
slender and short in the Carabidæ and Cicindelidæ, as well as those
insects whose food is liquid, such as Diptera. In the Lepidoptera it
varies in length, being in Sphinx quite long and bent into seven folds
(Fig. 309), while it is very short in the Psocidæ, Chrysomelidæ, and
Tenthredinidæ.

In the locust the ileum is traversed by six longitudinal folds with
intervening furrows; outside of each furrow is a longitudinal muscular
band. Seen from the inner surface the epithelium has an unusual
character, the cells in the middle of each of the flat folds being quite
large, polygonal in outline, while towards the furrows the cells become
very much smaller. The walls are double when seen in transverse section,
the inner layer consisting of epithelial cells resting on connective
tissue, the outer layer formed of circular muscles. The cuticula is
thin, but probably chitinous; it resembles that on the gastro-ileal
folds, except that there are no spinules, but unlike the cuticula of the
stomach it extends equally over the folds and the furrows. (Minot.) In
the cockroach the junction of the small intestine with the colon is
abrupt, a well-developed annular fold assuming the nature of a circular
valve. (Miall and Denny.)

=The gastro-ileal folds.=—In the locust the intestine is separated from
the chylific stomach by what Minot calls “the gastro-ileal folds,” which
form a peculiar valve. The urinary vessels open just underneath and in
front of this valve. In Melanoplus, and probably in the entire family of
Acrydiidæ, they are indicated as “dark spots, round in front and lying
at the anterior end of the ileum so as to form a ring around the
interior of the intestine.” They are 12 in number, and all alike. They
are pigmented and round in front where they are broadest and stand up
highest; they narrow down backwards, the pigment disappears, and they
gradually fade out into the ileal folds. Directly beneath them, and just
at the posterior end of the stomach, there is a strong band of circular
striated muscular fibres. The epithelium of these folds is covered with
minute conical spines, which are generally, but not always, wanting
between the folds. (Minot.)

=The colon.=—This section of the intestine (Fig. 319) is sometimes
regarded as a part of the rectum. In the locust the six longitudinal
folds of the ileum are continued into the colon, but their surface,
instead of being smooth as in the ileum, is thrown up into numerous
irregular curved and zigzag secondary folds. The cells of the epithelium
are of uniform size, and the layer is covered by a highly refringent
cuticula without spines; and, like that in the ileum, it rests on a
layer of connective tissue, beyond which follows (1) an internal coat of
longitudinal, and (2) an external coat of circular striated muscular
fibres. (Minot.)

[Illustration:

FIG. 319.—Digestive canal of _Lucanus cervus_: _G_, anterior muscles
of the pharynx; _H_, œsophagus; _I_, gizzard; _K_, chyle-stomach;
_L_, ilium; _M_, colon (cœcal part of); _N_, colon; _O_, rectum;
_a_, frontal ganglion of the vagus; _b_, vagus nerve; _c_, anterior
lateral ganglion connected with the vagus.—After Newport.
]

In butterflies (_Pontia brassicæ_), in _Sphinx ligustri_, and probably
in most Lepidoptera the colon is distinct from the rectum, and is
anteriorly developed into a very large more or less pyriform or
bladder-like cæcum (Figs. 309, 310), which in certain Coleoptera
(Dyticus, Fig. 320, _d_; Silpha, Necrophorus, etc.) is of remarkable
length and shape; it also occurs in Nepidæ (Fig. 327). In the cockroach
a lateral cæcum “is occasionally, but not constantly, present towards
its rectal end,” and a constriction divides the colon from the rectum.
(Miall and Denny.)

=The rectum.=—The terminal section of the hind-gut varies in length and
size, but is usually larger than the colon, and with thick, muscular
walls. In Lepidoptera it is narrow and short.

The rectum is remarkable for containing structures called rectal glands
(Fig. 298). Chun describes those of _Locusta viridissima_ as six flat
folds, formed by a high columnar epithelium and a distinct cuticula;
there is a coat of circular bands corresponding to the furrows between
the glands. Minot states that this description is applicable to the
locusts (Acrydiidæ) he has investigated, the only difference being in
the structural details of the single layers. He claims that the rectal
folds “do not offer the least appearance of glandular structure,”
neither is their function an absorbent one, as Chun supposed. From their
structure and position Fernald regards the rectal glands of Passalus as
acting like a valve, serving to retain the food in the absorptive
portions of the digestive tract till all nutriment is extracted.

[Illustration:

FIG. 320.—_Dyticus marginalis_, ♂ opened from the back: _a_, crop;
_b_, proventriculus; _c_, mid-intestine beset with fine cœcal
glands; _d_, long cœcal appendage of the colon; apodemes;
_B_{1}_-_B_{3}_, apodemes; _vhm_, coxal extensor muscle, moving the
hind leg; _ho_, testis; _dr_, accessory gland; _r_, penis; _e_,
reservoir of the secretion of the anal gland.—After Graber.
]

The epithelial folds of the larvæ of dragon-flies serve as organs of
respiration, the water being admitted into this cavity, and when
forcibly expelled serving to propel the creature forward. Paired and
single anal glands (repugnatorial) enter the rectum of certain
Coleoptera (Figs. 302, _l_; 317, _s_; 320, _e_).

=The vent (anus).=—The external opening of the rectum is situated in the
end of the body, in the vestigial 10th or 11th abdominal segment, and is
more or less eversible. It is protected above in caterpillars, and other
insects with 10 free abdominal segments, by the suranal plate. It is
bounded on the sides by the paranal lobes, while beneath is the
infra-anal lobe.

The anus is wanting in certain insects, and where this is the case the
hind-gut, owing to a retention of the embryonic condition, is usually
separated from the mid-intestine. (See p. 300.)

[Illustration:

FIG. 321.—Enteric canal of _Psyllopsis fraxinicola_: _œ_, œsophagus;
_md_, mid-intestine; _ed_, hind-intestine; _vm_, urinary vessels;
_s_, the coil formed by the hind-intestine and the most anterior
part of the mid-intestine.—After Witlaczil, from Lang.
]

Some remarkable features of the digestive canal in hemipterous
insects are noteworthy. In the Coccidæ, according to Mark, the
anterior end of the long mid-intestine forms, with the hinder end of
the œsophagus, a small loop, whose posterior end is firmly grown to
the wall of the rectum, and forms a cup-like invagination of the
latter. Then the rest of the tube-like stomach turns sidewise and
forms a large loop, which turns back on itself and occupies a large
part of the body-cavity. This loop receives on the anterior end,
near the œsophagus, the two urinary vessels, and forms just below
the opening into the rectum a short cæcum.

In other homopterous genera (Psyllidæ and some Cicadidæ) Witlaczil
describes nearly the same peculiarity, the mid-gut and part of the
intestine forming a loop growing together for a certain distance and
winding round each other (Fig. 321).

=Histology of the digestive canal.=—In all the divisions of the
digestive canal of insects the succession of the cellular layers
composing it is the same: 1st, a cuticula; 2d, an epithelium; 3d,
connective tissue; 4th, muscular tissue. In the locust, the first
division of the canal (fore-gut), there are two muscular coats, an
internal longitudinal and an external circular coat; the fibres are all
striated. The lining epithelium is not much developed, but forms a
thick, hard, and refringent cuticula, which is thrown up into spiny
ridges. In the second division (mid-gut, “stomach”) the epithelium is
composed of very high columnar cells, which make up the greater part of
the thickness of the walls, while the cuticula is very delicate,
slightly refringent, with no ridges, and is probably not chitinous; the
fibres of the muscular coats are not striated, while this division is
also distinguished by the presence of glandular follicles and folds. The
stomach and the cæcal appendages have all these peculiarities in common,
while no other part of the canal is thus characterized.

The third division (intestine and rectum) is composed of an epithelium,
the cells of which are intermediate in size between those of the fore
and mid gut. The cells are often pigmented, and they are covered by a
much thicker cuticula than that of the stomach, but which is not so
thick and hard as that of the œsophagus and proventriculus. The very
refringent cuticula is not thrown up into ridges, though in some parts
it is covered with delicate conical spines, which are very short. “The
epithelium and underlying connective tissue (_tunica propria_) are
thrown up into six folds, which run longitudinally, being regular in the
ileum and rectum (as the rectal glands), but very irregular in the
colon. Outside the depression between each two neighboring folds there
is a longitudinal muscular band, these making six bands. This peculiar
disposition of the longitudinal muscles does not occur in any other part
of the canal; it is, therefore, especially characteristic of the third
division.” (Minot.)

LITERATURE ON THE ORGANS OF DIGESTION

=Treviranus, G. R.= Resultate einiger Untersuchungen über den inneren
Bau der Insekten. (Verdauungsorgane von _Cimex rufipes_.) (Annal. d.
Wetterau. Gesells., 1809, i, pp. 169–177, 1 Taf.)

=Ramdohr, C. A.= Abhandlungen über die Verdauungswerkzeuge der
Insekten. 1811, vii, pp. 221, 30 Taf.

=Dutrochet, R. J. H.= Mémoire sur les métamorphoses du canal
alimentaire dans les insectes. (Journal de Physique, 1818, lxxxvi,
pp. 130–135, 189–204; Meckel’s Archiv, 1818, iv, pp. 285–293.)

=Suckow, F. W. L.= Verdauungsorgane der Insekten. (Heusinger’s
Zeitschr. f. organ. Physik., 1828, iii, pp. 1–89.)

=Doyère, L.= Note sur le tube digestif des Cigales. (Ann. Sc. nat.
Zool., 1839, 2 Sér., xi, pp. 81–85.)

=Grube, A. E.= Fehlt den Wespen- oder Hornissenlarven ein After oder
nicht? 1 Taf. (Müller’s Archiv für Physiol., 1849, pp. 47–74.)

=Sirodot.= Recherches sur les sécrétions chez les insectes. (Ann. Sc.
nat. Zool., 1858, 4 Sér., x, pp. 141–189, 251–328, 12 Pls.)

=Milne-Edwards, H.= Leçons sur la physiologie et l’anatomie comparée,
v, 1859, pp. 498–536, 581–638.

=Dufour, L.= Recherches anatomiques sur les Carabiques et sur
plusieurs autres insectes Coléoptères. Appareil digestif. (Ann. Sc.
nat., ii, 1824, pp. 462–482, 2 Pls.; iii, 1824, pp. 215–242, 5 Pls.,
pp. 476–491, 3 Pls.; iv, 1824, pp. 103–125, 4 Pls.; iv, 1825, pp.
265–283.)

—— Recherches anatomiques sur l’Hippobosque des chevaux. (Ann. Sc.
nat., 1825, vi, pp. 299–322, 1 Pl.)

—— Description et figure de l’appareil digestif de l’_Anobium
striatum_. (Ibid., xiv, 1828, pp. 219–222, 1 Pl.)

—— Recherches anatomiques sur les Labidoures. Appareil de la
digestion. (Ibid., xiii, 1828, pp. 348–354, 2 Pls.)

—— Recherches anatomiques et considerations entomologiques sur
quelques insectes Coléoptères, compris dans les familles des
Dermestins, des Byrrhiens, des Acanthopodes et des Leptodactyles.
Appareil digestif. (Ibid., Sér. 2, Zool., i, 1834, pp. 67–76, 2
Pls.)

—— Résumé des recherches anatomiques et physiologiques sur les
Hémiptères. (Ibid., pp. 232–239.)

—— Mémoire sur les métamorphoses et l’anatomie de la _Pyrochroa
coccinea_. Appareil digestif. (Ibid., Sér. 2, Zool., xiii, 1840, pp.
328–330, 334–337, 2 Pls.)

—— Histoire comparative des metamorphoses et de l’anatomie des
_Cetonia aurata_ et _Dorcus parallelepipedus_. Appareil digestif.
(Ibid., Sér. 2, Zool., 1824, xviii, pp. 174–176, 2 Pls.)

—— Anatomie générale des Diptères. Appareil digestif. (Ibid., Sér. 3,
Zool., i, 1814, pp. 248, 249.)

—— Histoire des métamorphoses et de l’anatomie du _Piophila
petasionis_. Appareil digestif. (Ibid., Sér. 3, Zool., i, 1844, pp.
372–377, 1 Pl.)

—— Études anatomiques et physiologiques sur les insectes Diptères de
la famille des Pupipares. Appareil digestif. (Ibid., Sér. 3, Zool.,
iii, 1845, pp. 67–73, 1 Pl.)

—— Recherches sur l’anatomie et l’histoire naturelle de l’_Osmylus
maculatus_. Appareil digestif. (Ibid., Sér. 3, Zool., ix, 1848, pp.
346–349, 1 Pl.)

—— Études anatomiques et physiologiques, et observations sur les
larves des Libellules. Appareil digestif. (Ibid., Sér. 3, Zool.,
xvii, 1852, pp. 101–108, 1 Pl.)

—— Recherches anatomiques sur les Hyménoptères de la famille des
Urocerates. Appareil digestif. (Ibid., Sér. 4, Zool., i, 1854, pp.
212–216, 1 Pl.)

—— Fragments d’anatomie entomologique. Sur l’appareil digestif du
_Nemoptera lusitanica_. (Ibid., Sér. 4, viii, 1857, pp. 6–9, 1 Pl.)

—— Recherches anatomique et considerations entomologiques sur les
Hémiptères du genre Leptopus. Appareil digestif. (Ibid., Sér. 4,
Zool., 1858, x, pp. 352–356, 1 Pl.)

—— Recherches anatomiques sur l’_Ascalaphus meridionalis_. Appareil
digestif. (Ibid., Sér. 4, xiii, 1860, pp. 200–202, 1 Pl.)

=Leydig, F.= Zur Anatomie von _Coccus hesperidum_. (Zeitschr. f.
wissens. Zool., v, 1853, pp. 1–12, 1 Taf.)

=Lubbock, J.= On the digestive and nervous System of _Coccus
hesperidum_. (Proc. Roy. Soc., ix, 1886, pp. 480–486; also Ann. Mag.
Nat. Hist., 1859, Ser. 3, iii, pp. 306–311.)

=Scheiber, S. H.= Vergleichende Anatomie und Physiologie der
Œstriden-Larven. V. Das chylo- und uropœtische System. (Sitzber. d.
k. Akad. d. Wissens. Wien. Math.-naturwiss. Cl., 1862, xlv, pp.
39–64, 1 Taf.)

=Gerstaecker, A.= Bronn’s Klassen und Ordnungen des Tierreichs. V.
Gliederfüssler. (Ernährungsorgane, pp. 87–105.)

=Graber, V.= Zur naheren Kenntnis des Proventriculus und der
Appendices ventriculares bei den Grillen und Laubheuschrecken.
(Sitzber. d. k. Akad. d. Wissensch. Wien. Mathem.-naturwiss. Cl.,
lix, 1869, pp. 29–46, 3 Taf.)

—— Ueber die Ernährungsorgane der Insekten und nächstverwandten
Gliederfüssler. (Mitteil. d. naturwiss. Vereins für Steiermark.
Graz, 1871, ii, pp. 181, 182.)

—— Verdauungssystem des Prachtkäfers. (Ibid., Graz, 1875.)

—— Die Insekten., i, 1877. (Verdauungsapparat, pp. 308–328.)

=Wilde, K. F.= Untersuchungen über den Kaumagen der Orthopteren.
(Archiv f. Naturgesch., xliii Jahrg., 1877, pp. 135–172, 3 Taf.)

=Simroth, H.= Ueber den Darmkanal der Larven von _Osmoderma eremita_
mit seinen Anhängen. (Giebel’s Zeitschr. f. d. ges. Naturwiss.,
1878, li, pp. 493–518, 3 Taf.)

=Müller, H.= Ueber die angebliche Afterlösigkeit der Bienenlarven.
(Zool. Anzeiger, 1881, pp. 530, 531.)

=Schiemenz, Paulus.= Ueber das Herkommen des Futtersaftes und die
Speicheldrüsen der Bienen, nebst einem Anhänge über das Riechorgan.
(Zeitschr. f. wissens. Zool., xxxviii, 1883, pp. 71–135, 3 Taf.)

=Rovelli, G.= Alcune ricerche sul tubo digerente degli Atteri,
Ortotteri e Pseudo-Neurotteri. (Como, 1884, p. 15.)

=Beauregard, H.= Structure de l’appareil digestif des Insectes de la
tribu des Vésicants. (Compt. rend. Acad. Paris, 1884, xcix, pp.
1083–1086.)

—— Recherches sur les Insectes vésicants., 1 Part, Anatomie. (Journ.
Anat. Phys. Paris, 1885, xxi Année, pp. 483–524, 4 Pls.; 1886, xxii
Année, pp. 85–108, 242–284, 5 Pls.)

—— Les Insectes vésicants, Paris, 1890, Chap. III, Appareil digestif,
pp. 63–99; (Phénomènes digestifs, pp. 161–170; Pls. 6–9.)

=Wertheimer, L.= Sur la structure du tube digestif de l’_Oryctes
nasicornis_. (Compt. rend. Soc. Biol. Paris, 1887, Sér. 8, iv, pp.
531, 532.)

=Kowalevsky, A.= Beitrage zur Kenntniss der nachembryonal Entwicklung
der Musciden. (Zeitschr. f. wissens. Zool., xlv, 1887, pp. 542–594,
5 Taf.)

=Schneider, A.= Ueber den Darm der Arthropoden, besonders der
Insekten. (Zool. Anzeiger, 1887, x Jahrg., pp. 139, 140.)

—— Ueber den Darmkanal der Arthropoden. (Zool. Beiträge von A.
Schneider, ii, 1887, pp. 82–96, 3 Taf.)

=Fritze, A.= Ueber den Darmkanal der Ephemeriden. (Berichte der
Naturforsch.-Gesellsch. zu Freiburg i. Br., 1888, iv, pp. 59–82, 2
Taf.)

=Emery, C.= Ueber den sogenannten Kaumagen einiger Ameisen. (Zeitschr.
f. wissens. Zool., 1888, xlvi, pp. 378–412, 3 Taf.)

=Meinert, F.= Contribution à l’anatomie des Fourmilions. (Overs.
Danske Vidensk. Selsk. Forhandl. Kjöbenhavn, 1889, pp. 43–66, 2
Pls.)

=Mingazzini, P.= Richerche sul canale digerente dei Lamellicorni
fitofage (Larve e Insetti perfetti). (Mitteil. Zool. Station zu
Neapel, ix, 1889–1891, pp. 1–112, 266–304, 7 Pls.)

=Fernald, Henry T.= Rectal glands in Coleoptera. (Amer. Naturalist,
xxiv, pp. 100, 101, Jan., 1890.)

=Visart, O.= Digestive canal of Orthoptera. (Atti Soc. Toscana Scient.
Natur., vii, 1891, pp. 277–285.)

=Eberli, J.= Untersuchungen an Verdauungstrakten von _Gryllotalpa
vulgaris_. (Vierteljahresschr. d. Naturforsch. Gesells. Zurich,
1892, Sep., p. 46, Fig.)

=Holmgren, Emil.= Histologiska studier öfver några lepidopterlarvers
digestionskanal och en del af deras Körtelartade bildningar. (Ent.
Tidskr. Årg. xiii, pp. 129–170, 1892, 6 Pls.)

=Ris, F.= Untersuchung über die Gestalt des Kaumagens bei den Libellen
und ihren Larven. (Zool. Jahrb. Abth. Syst., ix, 1896, pp. 596–624,
13 Figs.)

See also the works of Straus-Dürckheim, Newport, Mark, Witlaczil,
Vayssière, Landois, Jordan, Oudemans, Berlese, List, Grassi, Verson,
Miall and Denny, Leidy, Cheshire, Kowalevsky, Gehuchten, Locy, etc.

_b_. Digestion in insects

For the most complete and reliable investigation of the process of
digestion, we are indebted to Plateau, whose results we give, besides
the conclusions of later authors:

In mandibulate or biting insects, the food is conducted through the
œsophagus by means of the muscular coating of this part of the digestive
canal. Suctorial insects draw in their liquid food by the contractions
followed by the dilatations of the mid-intestine (chylific stomach).
Dragon-flies, Orthoptera, and Lepidoptera swallow some air with their
food.

Where the salivary glands are present, the neutral alkaline fluid
secreted by them has the same property as the salivary fluid of
vertebrates of rapidly transforming starchy foods into soluble and
assimilable glucose. In such forms as have no salivary glands, their
place is almost always supplied by an epithelial lining of the
œsophagus, or, as in the Hydrophilidæ, a fluid is secreted which has the
same function as the true salivary fluid.

Nagel states that the saliva of the larva of Dyticus is powerfully
digestive, and has a marked poisonous action, killing other insects, and
even tadpoles of twice the size of the attacking larva, very rapidly.
The larvæ not only suck the blood of their victims, but absorb the
proteid substances. Drops of salivary juice seem to paralyze the victim,
and to ferment the proteids. The secretion is neutral, the digestion
tryptic. Similar extra-oral digestion seems to occur in larvæ of
ant-lions, etc. (Biol. Centralbl., xvi, 1896, pp. 51–57, 103–112; Journ.
Roy. Micr. Soc., 1896, p. 184.)

In carnivorous insects and in Orthoptera, the œsophagus dilates into a
crop (ingluvies) ended by a narrow, valvular apparatus (or gizzard of
authors). The food, more or less divided by the jaws, accumulates in the
crop, which is very distensible; and, when the food is penetrated by the
neutral or alkaline liquid, there undergoes an evident digestive action
resulting, in carnivorous insects, in the transformation of albuminoid
substances into soluble and assimilable matter analogous to peptones,
and, in herbivorous insects, an abundant production of sugar from
starch. This digestion in the crop, a food-reservoir, is very slow, and,
until it is ended, the rest of the digestive canal remains empty.

“Any decided acidity found in the crop is due to the injection of acid
food; but a very faint acidity may occur, which results from the
presence in the crop of a fluid secreted by the cæcal diverticula of the
mesenteron.” (Miall and Denny.)

When digestion in the crop is accomplished, the matters are subjected to
an energetic pressure of the walls through peristaltic contractions, and
then, guided by the furrows and chitinous teeth, pass along or gradually
filter through the valvular apparatus or proventriculus, whose function
is that of a strainer.

At the beginning of the “chyle-stomach” (mesenteron) of Orthoptera are
glandular cæca which secrete a feebly acid fluid. This fluid emulsifies
fats, and converts albuminoids into peptones. It passes forwards into
the crop, and there acts upon the food.

In the mesenteron (mid-intestine) the food is acted upon by an alkaline
or neutral fluid, never acid, either secreted, as in Orthoptera, by
local special glands, or by a multitude of minute glandular cæca, as in
many Coleoptera, or by a simple epithelial layer. It has no analogy with
the gastric juices of vertebrates; its function differs in insects of
different groups; in carnivorous Coleoptera it actively emulsionizes
greasy matters; in the Hydrophilidæ it continues the process of
transformation of starch into glucose, begun in the œsophagus. In the
Scarabæidæ, it also produces glucose, but this action is local, not
occurring elsewhere; in caterpillars, it causes a production of glucose,
and transforms the albuminoids into soluble and assimilable bodies
analogous to peptones, and also emulsionizes greasy matters. Finally, in
the herbivorous Orthoptera there does not seem to be any formation of
sugar in the stomach itself, the production of glucose being confined to
the crop (jabot).

When digestion in the crop is finished, the proventriculus relaxes, and
the contents of the crop, now in a semi-fluid condition, guided by the
furrows and teeth, passes into the mesenteron, which is without a
chitinous lining, and is thus fitted for absorption.

The contents of the mid-intestine (chylific stomach) then slowly and
gradually pass into the intestine, the first anterior portion of which,
usually long and slender, is the seat of an active absorption. The
epithelial lining observed in certain insects seems, however, to
indicate that secondary digestion takes place in this section. The
reaction of the contents is neutral or alkaline.

The second and larger division of the intestine only acts as a stercoral
reservoir. (The voluminous cæcum occurring in Dyticidæ, Nepa, and
Ranatra, whether full or empty, never contains gas, and it is not, as
some have supposed, a swimming-bladder.) The liquid product secreted by
the Malpighian tubes accumulates in this division, and, under certain
circumstances, very large calculi are often formed. In his subsequent
paper on the digestion of the cockroach, Plateau states that in the
intestine are united the residue of the work of digestion and the
secretion of the urinary or Malpighian tubes, this secretion being
purely urinary.

These organs are exclusively depuratory and urinary, freeing the body
from waste products of the organic elements. The liquid they secrete
contains urea (?), uric acid and abundant urates, hippuric acid (?),
chloride of sodium, phosphates, carbonate of lime, oxalate of lime in
quantity, leucine, and coloring-matters.

The products of the rectal or anal glands vary much in different groups,
but they take no part in digestion, nor are they depuratory in their
nature.

Insects have nothing resembling chylific substances.[51] The products of
digestion, dissolved salts, peptones, sugar in solution, emulsionized
greasy matters, pass through the relatively delicate walls of the
digestive canal by osmose, and mingle outside of the canal with the
blood.

Whatever substances remain undigested are expelled with the excrements;
such are the chitin of the integuments of insects, vegetable cellulose,
and chlorophyll, which is detected by the microspectroscope all along
the digestive canal of phytophagous insects.

In his experiments in feeding the larvæ of Musca with lacmus, Kowalevsky
found that the œsophagus, food-reservoir, and proventriculus, with its
cæcal appendages, always remained blue, and had an alkaline reaction;
the mid-intestine, also, in its anterior portion, remained blue, but a
portion of its posterior half became deep red, and also exhibited a
strong reaction. The hind-intestine, however, always remained blue, and
also had an alkaline reaction. (Biol. Centralbl., ix, 1889, p. 46.)

=The mechanism of secretion.=—Gehuchten describes the process of
secretion in insects, the following extract being taken from his
researches on the digestive apparatus of the larva of Ptychoptera. The
products of secretion poured into the alimentary canal are more or less
fluid; for this reason, it is impossible to say when an epithelial cell
at rest contains these products. For the secreting nature of these cells
is only apparent at the moment when they are ready for excretion; then
the cellular membrane swells out, and a part of the protoplasmic body
projects into the intestinal cavity.

Before going farther, the terms _secretion_ and _excretion_ should, he
says, be defined. With Ranvier, he believes that the elaboration in the
protoplasm of a definite fluid substance is, _par excellence_, the
secretory act, while the removal of this substance is the act of
excretion.

[Illustration:

FIG. 322.—Different phases of the mechanism of secretion and of
excretion.—After Gehuchten.
]

A glandular cell of the chylific stomach, when at rest, is always
furnished with a striated “platform,” or flat surface, or face, on the
side facing the cavity of the stomach, and the free edge of the
platform, or plateau, is provided with filaments projecting into the
digestive cavity (Fig. 322, _f_). These glandular cells, when active,
differ much in appearance. In a great number, the platform (plateau) has
disappeared, and is replaced by a simple, regular membrane. During the
process of secretion, a finely granular mass, in direct continuity with
the protoplasm, swells, and raises the membrane over the entire breadth
of the cell, causing it to project into the intestinal cavity (Fig. 322,
_A_, _B_). These vesicles, or drops of the secretion, whether free or
still attached by a web to the cells, are clear and transparent in the
living insect, but granular in the portions of the digestive canal fixed
for cutting into sections. Gehuchten then asks: “How does a cell gorged
with the products of secretion empty itself?” Both Ranvier and also
Heidenhain believe that one and the same glandular cell may secrete and
excrete several times without undergoing destruction, but their
researches made on salivary glands have not answered the question.
Gehuchten explains the process thus: when the epithelial cell begins to
secrete, the clear fluid elaborated in the protoplasm of the cell
increases the intracellular tension, until, finally, the fluid breaks
through certain weak places in the swollen basal membrane of the
platform, and then easily passes through the closely crowded filaments,
and projects out into the intestinal cavity as a pear-shaped vesicle of
a liquid rich in albumens at first attached to the free face of the
cell, but finally becoming free, as at Fig. 322, _A_, _B_.

When the elaboration of the substance to be secreted is more active, the
mechanism of the secretion is modified. The basal membrane of the
platform may then be raised at several places at once; instead of a
single vesicle projecting into the intestinal cavity, each cell may
present a great number more or less voluminous. If all remain small and
rapidly detach themselves from the glandular cell, the filaments of the
platform are simply separated from each other at different points of the
free face, as in Fig. 322, _C_. On the other hand, when the different
vesicles of a single cell become larger, the filaments of the platform
are compressed and crowded against each other in the spaces between the
vesicles remaining free, and the undisturbed portions of the platform
appear homogeneous (Fig. 322, _D_). After the excretion of the secretory
products by this process of strangulation, the cell then assumes the
aspect of a glandular cell at rest, and may begin again to form a new
secretion.

To sum up: The process of excretion may occur in two ways:

1. Where the membrane ruptures and the substances secreted are sent
directly out into the digestive cavity. 2. Where the vesicles become
free by strangulation, floating in the glandular or intestinal cavity,
and ending by rupturing and coming into contact with the neighboring
vesicles or with the food.

=Absorbent cells.=—Besides the glandular or secreting cells in
Ptychoptera, there is between the two regions of the chyle-stomach lined
with these cells a region about a centimetre long composed of absorbent
cells. The absorbent cells are very large, polygonal, and contain a
large nucleus, in which is a striated convoluted chromatic cord.

The food on entering the chyle-stomach is brought into contact with the
products secreted in the proventriculus, in the first part of the
chyle-stomach, and in the tubular glands. These products of secretion
act on the food, extracting from them useful substances which they
render soluble. These substances, after having been absorbed by the
absorbent cells in the middle region of the stomach, undergo special
modifications, and are transformed into solid products, which are
situated at the bottom of these cells. Afterwards the alimentary
substances freed from a portion of their useful substances are again
placed in contact with the products of secretion in the distal part of
the chylific ventricle, and reach the terminal part of the intestine.

“The products of secretion,” adds Gehuchten, “diverted into the
intestinal canal do not come into immediate contact with the alimentary
substances; they are separated from it by a continuous, structureless,
quite thick membrane (the peritrophic membrane), which directly envelops
the cylinder of food matters, extending from the orifice of the
œsophageal valvule to the end of the intestine. Between this membrane
and the free face of the epithelial lining there exists a circular
space, into which are thrown and accumulate the excreted substances. The
latter then cannot directly mingle with the aliments; but when they are
liquid they undoubtedly pass through this membrane by osmose, and thus
come into contact with the nutritive substances. It is the same with the
products of absorption. The absorption of soluble products of the
intestinal cavity is not then so simple a phenomenon as it was at first
thought to be, since these products are nowhere brought into immediate
contact with the absorbent cells” (pp. 90, 91).

The most recent authority, Cuénot, states that absorption of the
products of digestion takes place entirely in the mid-intestine, and in
its cæca when these are present. The mid-intestine exercises a selective
action on the constituents of the food comparable to the action of the
vertebrate liver.

LITERATURE ON THE PHYSIOLOGY OF DIGESTION

=Davy, J.= Note on the excrements of certain insects, and on the
urinary excrement of insects. (Edinburgh New Phil. Journ., 1846, xl,
pp. 231–234, 335–340; 1848, xlv, pp. 17–29.)

—— Some observations on the excrements of insects, in a letter
addressed to W. Spence. (Trans. Ent. Soc. London, Ser. 2, iii, 1854,
pp. 18–32.)

=Bouchardat, A.= De la digestion chez le ver à soie. (Revue et Mag. de
Zool., Sér. 2, 1851, iii, pp. 34–40.)

=Lacaze-Duthiers, H., et A. Riche.= Mémoire sur l’alimentation de
quelques insects gallicoles et sur la production de la graisse.
(Ann. Scienc. natur., 1854, Sér. 4, ii, pp. 81–105.)

=Basch, S.= Untersuchungen über das chylopoetische und uropoetische
System von _Blatta orientalis_. (Sitzungsber. d. math.-naturwiss.
Classe d. Akad. d. Wissensch. Wien., 1858, xxxiii, pp. 234–260, 5
Taf.)

=Lambrecht, A.= Der Verdauungsprozess der stickstoffreichen
Nährmittel, welche unsere Bienen geniessen, in den dazu geschaffenen
Organen derselben. (Bienenwirtschaftl. Centralbl., viii Jahrg.,
1872, pp. 73–78, 83–89.)

=Plateau, F.= Recherches sur les phénomènes de la digestion chez les
insectes, (Mém. Acad. roy. de Belgique, Sér. 2, xli, 1 Part, 1873,
pp. 124, 3 Pls.)

—— Note additionelle au mémoire sur les phénomènes de la digestion
chez les insectes. (Bull. Acad. roy. de Belgique, Sér. 2, xliv,
1877, pp. 710–733.)

=Tursini, G. Fr.= Un primo passo nella ricerca dell’ assorbimento
intestinale degli artropodi. (Rend. d. R. Accadem. di Sc. fis. e
matemat. di Napoli, 1877, xvi, pp. 95–99, 1 Pl.)

=Jousset de Bellesme=, Physiologie comparée. Recherches expérimentales
sur la digestion des insectes et en particulier de la blatte. Paris,
1876, vii, and 96 pp., 3 Pls.

—— Recherches sur les fonctions des glandes de l’appareil digestif des
Insectes. (Compt. rend., lxxxii, Paris, 1876, pp. 97–99.)

—— Travaux originaux de physiologie comparée. (i, Insectes, Digestion,
Métamorphoses.) Paris, 1878, 5 Pls.

=Simroth, H.= Einige Bemerkungen über die Verdauung der Kerfe.
(Zeitschr. f. d. gesammten Naturwiss., xli, 1878, pp. 826–831.)

=Krukenberg, C. Fr. W.= Versuche zur vergleichenden Physiologie der
Verdauung und vergleichende physiologische Beiträge zur Kenntnis der
Verdauungsvorgange. (Untersuch, a. d. physiolog. Institut d.
Universität Heidelberg, 1880, i, 4, pp 327, Figs.; ii, 1, p. 1,
Figs.)

=Metschnikoff, E.= Untersuchungen über die intrazelluläre Verdauung
bei wirbellosen Tieren. (Arb. d. zool. Instit. Wien., 1883, v, pp.
141–168, 2 Taf.)

=Locy, William A.= Anatomy and physiology of the family Nepidæ. (Amer.
Naturalist, xviii, 1884, pp. 250–255, 353–367, 4 Pls.)

=Vangel, E.= Beiträge zur Anatomie, Histiologie, und Physiologie des
Verdauungsapparates des Wasserkäfers, _Hydrophilus piceus_.
(Termész. Füzet., x, 1886, pp. 111–126 (in Hungarian); pp. 190–208
(in German), 1 Pl.)

=Schönfeld.= Die physiologische Bedeutung des Magenmundes der
Hönigbiene. (Archiv f. Anat. u. Physiol., Physiol. Abt., 1886, pp.
451–458.)

=Faussek, V.= Beiträge zur Histiologie des Darmkanals der Insekten.
(Zeitschr. f. wiss. Zool., 1887, xl, pp. 694–712, 1 Taf.; Abstract
in Zool. Anz., Jahrg. x, pp. 322, 323, 1 Taf.)

=Frenzel, J.= Ueber Bau und Thätigkeit des Verdauungskanals der Larve
des _Tenebrio molitor_, mit Berücksichtigung anderer Arthropoden
(Berlin. Ent. Zeitschr., 1882, pp. 267–316, 1 Taf.); Inaug.-Diss.
Göttingen, 1882.

—— Einiges über den Mitteldarm der Insekten, sowie über
Epithel-regeneration. (Archiv f. Mikrosk. Anat., 1885, xxvi, pp.
229–306, 3 Taf.)

—— Zum feineren Bau des Wimperapparates. (Ibid., 1886, xxviii, pp.
53–80, 1 Taf.)

—— Die Verdauung lebenden Gewebes und die Darmparasiten. (Archiv f.
Anat., 1891.)

=Gehuchten, A. van.= Recherches histologiques sur l’appareil digestif
de la _Ptychoptera contaminata_, I Part. Étude du revêtment
épithélial et recherches sur la sécrétion. (La Cellule, 1890, vi,
pp. 183–291, 6 Pls.)

=Cuénot, L.= Études physiologiques sur les Orthoptères. (Arch. Biol.,
xiv, 1895, pp. 293–341, 2 Pls.)

=Needham, James G.= The digestive epithelium of dragon-fly nymphs.
(Zool. Bull., i, 1897, Chicago, pp. 104–113, 10 Figs.)

With the writings of Mingazzini (see p. 323), Kowalevsky, Ranvier,
Haidenhain, Beauregard (p. 323), Sadones.

THE GLANDULAR AND EXCRETORY APPENDAGES OF THE DIGESTIVE CANAL

Into each primary division of the digestive canal open important glands.
The salivary and silk-glands are offshoots of the œsophagus (stomodæum);
the cœcal appendages open into the stomach (mesenteron), while the
urinary tubes grow out in embryonic life from the primitive intestine
(proctodæum), and there are other small glands which are connected with
the end of the hind-intestine.

_a._ The salivary glands

We will begin our account of these glands with those of the Orthoptera,
where they are well developed. In the cockroach a large salivary gland
and accompanying reservoir lie on each side of the œsophagus and crop.
The gland is a thin, leaf-like, lobulated mass, divided into two
principal lobes. These open into a common trunk, which after receiving a
branch from a small accessory lobe, and from the salivary reservoir,
unites with its fellow to form the unpaired salivary duct which opens
into the under side of the lingua. Each salivary reservoir is a large
oval sac with transparent walls. (Miall and Denny, also Figs. 299, _sr_,
and 327.) The ducts and reservoirs have a chitinous lining, and the
ducts are, like the tracheæ, surrounded by a so-called spiral thread, or
by separate, incomplete, hooplike bands, which serve to keep the duct
permanently distended. In the locust (Fig. 298) the lobules are more
scattered, forming small separate groups of acinose glands. In the
embryo of Forficula Heymons has observed a pair of salivary glands
opening on the inner angle of the mandibles, a second pair opening in
the second maxillæ, while a third pair of glands, whose function is
doubtful, is situated in the hinder part of the head, opening to the
right and left on the chitinous plate (postgula) behind the submentum.
In Perla, there are two pairs segmentally arranged (Fig. 343).

[Illustration:

FIG. 323.—Left side of the head of the silkworm: _a_, adductor muscle
of the mandible, from which the muscular fibres have been removed;
_b_, upper fibres of the same; _c_, lower fibres cut away to show
the adductor muscle (_e_); _d_, fibres inserted on the accessory
adductor lamella; _f_, œsophagus, much swollen; _g_, salivary gland;
_h_, dorsal vessel; _i_, _l_, tracheæ of the mandibular muscles;
_k_, trachea; _n_, optic nerve.—After Blanc.
]

[Illustration:

FIG. 324.—Lower side of the head of the silkworm exposed, the spinning
apparatus, the œsophageal ganglion, and the adductor of the left
mandible removed: _M_, mandible; _P_, abductor of the mandible; _R_,
adductor; _N_, salivary gland attached at _O_ to the edge of the
adductor muscle; _o_, _o_, transverse portion of the “hyoid”; 3,
masticator nerve and its recurrent branch (7); _L_, tongue cut
horizontally.—After Blanc.
]

Here we might refer to a pair of glands regarded by Blanc as the true
salivary glands. They do not appear to be the homologues of the salivary
glands of other insects, though probably functioning as such. The
functional salivary glands of lepidopterous larvæ have been overlooked
by most entomotomists, and the spinning glands have been, it seems to
us, correctly supposed to be modified salivary glands. Lucas also
regards those of case-worms (Trichoptera) as morphologically salivary
glands. Those of the silkworm were figured by Réaumur (Tom. i, Pl. v,
Fig. 1), but not described; while those of Cossus, which are voluminous,
were regarded by Lyonet as “_vaisseaux dissolvans_.” Dr. Auzoux (1849),
in his celebrated model of the silkworm, represented them accurately,
while Cornalia briefly described them as opening into the mouth. The
first satisfactory description is that of Blanc (1891), who states that
in the silkworm “the two salivary glands” are small, flexuous, yellow
tubes, which occupy a variable position on the sides of the œsophagus
(Fig. 323). The glandular portion passes into the head, ending at the
level of the adductor plate of the mandibles (Fig. 324, _o_), and
entering the buccal cavity at the base of the mandible, as seen in Fig.
323. It is plain, when we recognize the direct homology of the
silk-glands of the caterpillars with the salivary glands of other
insects, and of the spinneret with the hypopharynx, that these so-called
“salivary glands” in lepidopterous larvæ are different structures. They
are probably modified coxal glands, belonging to the mandibular segment.

[Illustration:

FIG. 325.—One of the two salivary glands of _Cæcilius burmeisteri_:
_d_, excretory duct; _cn_, the lumen or canal; _cg_, gland-cells;
_ct_, salivary fluid.—After Kolbe.
]

The polygonal epithelial cells of these glands contain branched
nuclei, recalling those of the spinning-glands. In those
caterpillars which feed on leaves, the salivary glands are slightly
developed, but in such as bore into and eat wood, as the Cossidæ,
the glands are, as figured by Lyonet, very large, forming two
sausage-shaped bodies passing back to the beginning of the
mid-intestine, each ending in a long convoluted filament. The
salivary glands of the imago are very long and convoluted (Fig. 310,
_sd_).

In the Panorpidæ these glands differ in the sexes, the males having
three pairs of very long tortuous tubes, while, in the females, they
are reduced to two indistinct vesicles. (Siebold.)

In the Diptera in general there are two pairs, one situated in the beak,
the other in the thorax. In the larvæ there is a single pair (Fig. 341).
Kraepelin describes a third pair in the Muscidæ at the point of
transition from the fulcrum to the œsophagus, but Knüppel has apparently
found only what may be fat cells at this point, so that the supposed
presence of a third pair in Diptera needs confirmation. In the Psocidæ
there are two salivary glands, of simple tubular shape (Fig. 325).

In the Nepidæ the salivary glands are four in number, and of
conglomerate structure, two being long and extending back into the
beginning of the abdomen, while the other two are about one-fourth as
long. (Figs. 327, 328.) In Cicada, besides a pair of simple tortuous
tubes, there is in the head another pair of glands, each composed of two
tufts of short lobes, situated one behind the other. (Dufour.) In many
Hemiptera (Pyrrhocoris, Capsus, etc.) there is but a single pair, each
gland consisting of four lobes; in the Coccidæ each gland is divided
into two lobes (Fig. 326); in the Aphidæ, according to Witlaczil, they
consist of two lobes grown together. In the Psyllidæ they are said to be
absent.

In _Phylloxera vastatrix_ the saliva is forced through a salivary
passage out of the duct and into the mouth by a pumping apparatus
furnished with special muscles. (Krassilstschik.)

In the Odonata acinose glands are present in the imago, but not in the
nymph until in its last stage, Poletaiew accounting for their absence in
the earlier stages by the fact that the larva swallows more or less
water while taking its food.

In the Coleoptera, as we have observed in Anopthalmus, there are three
pairs of salivary glands (Fig. 74). In the Blapsidæ these glands consist
of many ramifying tubes united on each side of the œsophagus into a
single duct; in others they are but slightly developed, while in still
others they are wanting.

The salivary glands are most highly differentiated in the Hymenoptera,
and especially in the bees (Bombus and Apis), where Schiemenz found not
less than five systems of glands (Fig. 329; also 87), of which four
systems are paired. One pair of these glands lies in the tongue, three
in the head, and one in the thorax.

[Illustration:

FIG. 326.—Acinous salivary glands of _Orthezia cataphracta_. In some
acini the nuclei and boundaries of the cells are shown.—After List,
from Field’s Hertwig.
]

System I is situated in the head, and consists of unicellular
glands; the duct from each cell leads into a common, strongly
chitinized duct, opening into the gullet.

System II, composed of acinose glands, lies also in the head; its
duct is united with that of System III, situated in the thorax.
(Fig. 329, 2, 3.)

System IV is situated at the base of the upper surface of the
mandibles, and forms a delicate sac lined within with glandular
cells; its duct opens at the insertion of the mandibles.

System V lies in the beak, and is a single gland consisting of
unicellular glands; it opens into the common opening of Systems II
and III. This system is wanting in the honey-bee, but occurs in
Bombus and other genera.

[Illustration:

FIG. 327.—Appendages of digestive canal of Belostoma.—After Locy.
]

[Illustration:

FIG. 328.—Salivary and other glands of Ranatra.—After Locy.
]

In all the five systems there constantly occur three cellular
layers: the intima, epithelial, and propria. As regards their origin
Schiemenz states that Systems I and IV are new structures, that
System III arises in part, and Systems II and V wholly, from the
silk-glands of the larva. As the glands differ much in the sexes,
and in different species and genera, Schiemenz believes that their
function is very manifold.

In addition to those previously discovered by Schiementz, Bordas has
detected two additional pairs of salivary glands in the worker and
male honey-bee, _i.e._ the internal mandibular and sublingual
glands, so that in Apis there are in all six pairs, and apparently
one unpaired.

The delicate chitinous external layer of the gland is perforated by many
very fine pores through which the salivary fluid secreted by the
epithelial cells passes into the salivary duct. The glands are
externally bathed by the blood.

In many insects, including lepidopterous larvæ, the single median
opening of the salivary duct is converted into a spraying apparatus.

In the adult Lepidoptera, according to Kirbach:—

[Illustration:

FIG. 329.—Salivary glands of the honey-bee: systems No. 1–3, × 15:
_sv_, salivary valve (of systems 2 and 3) at base of tongue; _lp_,
labial palpus; _mx_, maxilla; _so_, salivary opening of system 1 in
hypopharyngeal plate; _no_, openings in plate for termination of
taste-nerve; _œ_, œsophagus; _sd_, salivary duct; _b_, junction of
ducts of system No. 2; _c_, junction of ducts of system No. 3; _sc_,
_sc_, salivary sacs; _fl_, front lobe; _bl_, back lobe; _a_,
chitinous duct, with spiral thread. _B_, single acinus of system No.
1, × 70: _n_, nucleus; _st_, salivary tract; _d_, large duct. _C_,
single pouch, or acinus, from system No. 2: _a_, propria or outer
membrane; _sc_, secreting cells. _D_, termination of system No.
3:_{1},_{2},_{3},_{4}, lines marking end of section; _d_, duct in
section; _sc_, secreting cells in section; _n_, nucleus.—After
Cheshire.
]

“Its lower half forms a thick chitinous gutter, with a concave cover
above, in which the similarly shaped upper half lies encased, so
that between the two only a small semicircular opening remains.
Powerful muscles extend from the cover to the lower side and to the
two ridges of the bottom plate; through their contraction the upper
channel is elevated, and presses out of the hinder part of the ducts
into the space thus formed a great quantity of the saliva, which by
allowing the contraction of the cover-muscle through the
crevice-like opening, which is situated in the lower edge of the
mouth-opening, becomes squeezed out in order either to mix with the
fluid where the 2d maxillæ fuse, passing up into the canal in the
proboscis, or to penetrate into and thus dilute the semi-fluid or
solid substances taken, into the proboscis.”

The morphology and general relations of the salivary glands have
been sketched out by Hatschek, Patten, and by Lucas, from
observations on those of the case-worms or larval Trichoptera.

[Illustration:

FIG. 330.—Eight pairs of glands of Andrena: I, thoracic; II,
postcerebral; III, supracerebral; IV, lateropharyngeal; V,
mandibular; VI, internomandibular; VII, sublingual; VIII, lingual;
_Md_, mandible; _L_, tongue; _o_, eye; _œ_, œsophagus; _J_,
honey-sac.—After Bordas.
]

Patten states that the spinning-glands in Neophylax are formed by a
pair of ectodermal invaginations on the ventral side of the embryo,
between the base of the 2d maxillæ and the nervous cord. They
increase rapidly in length, and “they also unite to form a common
duct, which opens at the end of the upper lip.”

The salivary glands in the same insect are “formed by invagination
of the ectoderm on the inner sides of the mandibles, in the same
manner as are the spinning glands.”

Lucas has shown that in trichopterous larvæ (Anabolia) there are
three pairs of salivary glands in the head, which are serially
arranged. The first pair belong to the mandibular, the second pair
to the 1st maxillary, and the third pair, or spinning glands, to the
2d maxillary segment. The first or mandibular glands open into the
mouth at the base of the mandibles directly behind the dorsal
condyle. The second pair open between the 1st and 2d maxillæ; at the
base of the latter, near the ventral condyle of the mandibles. The
third pair open into the hypopharynx, which is modified to form the
spinneret. Lucas agrees with Korschelt in regarding them as modified
coxal glands, Schiemenz having previously regarded the headglands of
the imago of the bee as belonging to the segments bearing the three
pairs of buccal appendages, so that each segment originally
contained a pair of glands. It is thus proven that the silk-glands
are modified salivary glands adapted to the needs of spinning larvæ,
and indeed in the imago the sericteries revert to their primitive
shape and use as salivary glands.

The serial arrangement of the salivary glands in the Hymenoptera,
where the number varies from five to ten pairs, is clearly proved by
Bordas. He has detected five more pairs than were previously known,
and names the whole series as follows:—1, the thoracic salivary
glands, which are larger than the others, and nine other pairs,
which are all contained in the head as follows: 2, postcerebral; 3,
supracerebral; 4, lateropharyngeal; 5, mandibular; 6,
internomandibular, situated on the inner side of the base of
mandible; 7, sublingual; 8, lingual (these and 1 to 7 common to all
Hymenoptera); 9, paraglossal (in Vespidæ); 10, maxillary (very
distinct in most wasps). These glands do not all occur in the same
species, being more or less atrophied.

Bordas further shows the segmental arrangement of the cephalic
glands by stating that the supracerebral glands correspond to the
antennal segment, the sublingual glands to the labial, the
mandibular glands (external and internal) to the mandibular segment,
the maxillary glands to the 1st maxillary segment, the lingual
glands to the 2d maxillary segment, while the thoracic and
postcerebral salivary glands, he thinks, correspond to the ocular
segment, a view with which we are indisposed to agree, although
conceding that each of the six segments of the head has in it at
least one pair of salivary glands.

=Functions of the different salivary glands in Hymenoptera.=—The
secretion of the thoracic glands is feebly alkaline. The
postcerebral salivary glands, considered by Ramdohr to be organs of
smell, secrete, like the preceding, a distinctively alkaline fluid,
which mingles with the products of the thoracic glands. The
supracerebral glands, also equally well developed in all
Hymenoptera, though much atrophied in the females and especially the
males of _Apis mellifica_, also in the Vespinæ and Polistinæ,
secrete an abundant, feebly acid liquid, which is actively concerned
in digestion.

As to the mandibular glands, which Wolf supposed to be olfactory
organs, their acid secretion, though smelling strongly, acts
energetically on the food as soon as introduced into the mouth.

The sublingual glands, atrophied in most Apidæ, but relatively
voluminous in Sphegidæ, Vespinæ, Polistinæ, Crabronidæ, etc., empty
their secretion into a small prebuccal excavation, where accumulate
vegetable and earthy matters collected by the tongue, and the saliva
secreted by these glands, acts upon them before they pass into the
pharynx. The lingual glands secrete a thick, sticky liquid, which
causes foreign bodies to adhere to the tongue, and also agglutinates
alimentary substances. The uses of the other glands, maxillary and
paraglossal, are from their minuteness undetermined. (Bordas.)

LITERATURE ON THE SALIVARY GLANDS

=Leydig, F.= Zur Anatomie der Insekten. (Archiv Anat. und Phys. 1859.)

—— Untersuchungen zur Anatomie und Histiologie der Tiere. Bonn, 1883,
pp. 174, 8 Taf.

—— Intra- und interzellulare Gänge. (Biolog. Centralblatt, x, 1890,
pp. 392–396.)

=Dohrn, A.= Zur Anatomie der Hemipteren. (Stettin. Entom. Zeit., 1866,
salivary glands, pp. 328–332.)

=Kupffer, C.= Die Speicheldrüsen von _Periplaneta orientalis_ und ihr
Nervenapparat. (Beiträge zur Anatomie und Physiol., 1875.)

=Schiemenz, P.= Ueber das Herkommen des Futtersaftes und die
Speicheldrüsen der Biene. (Zeitschr. f. wissens. Zool., xxxviii,
1883, pp. 71–135, 3 Taf.)

=Korschelt, E.= Ueber die eigentümlichen Bildungen in den Zellkernen
der Speicheldrüsen von _Chironomus plumosus_. (Zool. Anzeiger, 1884,
pp. 189–194, 221–225, 241–246.)

=Hofer, B.= Untersuchungen über den Bau der Speicheldrüsen und des
dazu gehörenden Nervenapparates von Blatta. (Nova Acta d. Kais.
Leopold.-Carol. Deutsch. Akad. d. Naturforscher, li, 1887, pp.
345–395, 3 Taf.)

=Knüppel, A.= Ueber Speicheldrüsen von Insekten. (Archiv für Naturg.,
1887, Jahrg. 52, pp. 269–303, 2 Taf.)

=Blanc, Louis.= La tête du _Bombyx mori_ à l’état larvaire, anatomie
et physiologie. (Extrait des Travaux du Laboratoire d’Études de la
Soie, 1889–1890; Lyon, 1891, p. 180, many figs.)

=Bordas, L.= Anatomie des glandes salivaires des Hyménoptères de la
famille des Ichneumonidæ. (Zool. Anzeiger, 1894, pp. 131–133.)

—— Glandes salivaires des Apides, _Apis mellifica_. ♂ and ♀. (Comptes
rendus Acad. Sc., Paris, cxix, pp. 363, 483, 693–695, 1894; also two
articles in Bull. Soc. Philomath. Paris, 1894, pp. 5, 12, 66.)

—— Appareil glandulaire des Hyménoptères. (Ann. Sc. Nat. Zool., xix,
Paris, 1894, pp. 1–362, 11 Pls.) (See also p. 366.)

=Berlese, Antonio.= Le cocciniglie Italiane viventi sugli agrumi.
Firenze, 1896, 12 Pls. and 200 Figs.

With the writings of Mark, Minot, Locy, List, Krassilstschik, Nagel
(1896).

_b._ The silk or spinning glands, and the spinning apparatus

The larvæ of certain insects, chiefly those of the Lepidoptera, possess
a pair of silk or spinning glands (sericteries) which unite to form a
single duct opening in the upper lip at the end of the lingua, which is
modified to form the spinneret. (See pp. 71, 75.) All caterpillars
possess them, and they are best developed in the silkworms, which spin
the most complete cocoon. Silk-glands also occur in the larvæ of the
Tenthredinidæ, in the case-worms or larval Trichoptera, also in certain
chrysomelid beetles (Donacia, Hæmonia), and in a weevil (Hypera). In a
common caddis-worm (Limnophilus) the glands are of a beautiful pale
violet-blue tint, and two and a half times as long as the larva itself;
viz. the body is 20 mm. and the glands 55 mm. in length.

In caterpillars the glands are of tubular shape, shining white, and much
like the ordinary simple tubular salivary glands of the imago. When only
slightly longer than the body they are twice folded, the folds parallel
and situated partly beneath and partly on the side of the digestive
canal; not usually, when folded in their natural position, extending
much behind the end of the stomach; but in the silkworms they are so
long and folded as to envelop the hinder part of the canal. In geometrid
caterpillars the glands when stretched out only reach slightly beyond
the end of the body; in Datana they are half again as long as the body.
Helm thus gives their relative length in certain Eurasian caterpillars,
and we add that of _Telea polyphemus_:—

_Vanessa io_ length of body 32 mm.; of the silk glands 26 mm.
_Smerinthus tiliæ_ length of body 63 mm.; of the silk glands 205 mm.
_Bombyx mori_ length of body 56 mm.; of the silk glands 262 mm.
_Antheræa yamamaya_ length of body 100 mm.; of the silk glands 625 mm.
_Telea polyphemus_ length of body 60 mm.; of the silk glands 450 mm.

Thus in Telea the silk-glands are about 18.50 inches in length, being
about seven times as long as the body.

For the most complete accounts of the spinning glands of Lepidoptera and
their mechanism we are indebted to Helm and to Blanc, and for that of
the Trichoptera to Gilson.

The unpaired portion, or spinning apparatus (_filière_ of Lyonet), is
divided into two portions; the hinder half being the “thread-press,” the
anterior division the “directing tubes.” The silk material, stored up in
the thickest portion of the glands, passes into the thread-press (Fig.
334, _A_), which is provided with muscles which force the two double
ribbon-like threads through the directing tube, as wire is made by
molten iron being driven through an iron plate perforated with fine
holes. The entire spinning apparatus, or _filator_, as we may call it,
is situated in the tubular spinneret. The opening of the spinneret is
directed anteriorly, and the anterior end of the directing tube passes
directly into this opening so that the directing tube may be regarded as
an invagination of the lingua.

The silk thread which issues from the mouth of the spinneret is, as
Leeuwenhoek discovered, a double ribbon-like band, as may be seen in
examining the silk of any cocoon.

=The process of spinning.=—Since the appearance of Helm’s account,
Gilson, and also Blanc, have added to our knowledge of the way in which
the silk is spun and of the mechanism of the process. Gilson has
arrived, in regard to the function of the press or filator, at the
following conclusions: 1, the press regulates the thread, it compresses
it, gives it its flattened shape; 2, it regulates the layer of gum[52]
(grès) which surrounds the thread; 3, it may render the thread immovable
by compressing it as if held by pincers.

The process of spinning in the silkworm, says Blanc, comprises all the
phenomena by which the mass of silk contained in the reservoir is
transformed into the silk fluid of which the cocoon is spun. The
excretory canals each contain a cylindrical thread of silk having a mean
diameter of 0.2 mm. and surrounded by a layer of gum (_grès_) which in
the fresh living organ exactly fills the annular space situated between
the fibroin cylinder and the wall. Arrived within the common duct, the
two threads receive the secretion of Filippi’s gland, where the silken
fluid is formed, but has not yet assumed its definite external
characters. The two threads press through the common canal and arrive at
the infundibulum (Fig. 334, _c_) of the press, at the bottom of which is
situated the orifice of the spinning canal, almost completely divided
into two by the sharp edge of the rachis (Figs. 334, _a_, 335, _l_). The
threads each pass into one of the two grooves, and the layer of gum
(_grès_) fills the rest of the canal of the press or filator.

[Illustration:

FIG. 331.—Longitudinal section of the spinneret: _a_, horizontal
portion of the tongue; _b_, vertical portion; _c_, _f_, circle of
the tongue; _d_, tongue-pad; _e_, orifice of the spinneret; _g_,
body of the lyre; _h_, prebasilar membrane forming a fold; _i_,
internal canal of the spinneret; _k_, filator.—After Blanc.
]

[Illustration:

FIG. 332.—The lower lip (labium) of _Bombyx mori_, isolated, seen from
the left side: _A_, lyre; _B_, spinneret; _C_, labial palpus; _D_,
vertical part of the labium; _E_, horizontal part of the same; _H_,
_L_, silk-canal; _K_, right gland of Filippi; _L_, canal of the left
gland; _N_, labial nerve; _a_, oblique fibre of the elevator of the
labium; _b_, right fibre of the same; _c_, depressor of the labium;
_d_, superior spinning muscles.
]

[Illustration:

FIG. 333.—The labium in a horizontal position, seen from the side:
_f_, the filator or press situated under the external part of the
spinneret (_d_), between the branches (_b_), of the lyre (_a_); _e_,
labial palp; _c_, tongue.
]

[Illustration:

FIG. 334.—Longitudinal section of the spinneret and press (_filator_):
_A_, filator or press; _B_, spinneret; _C_–_D_, body of the lyre;
_F_, lower part of the labium; _E_, common canal; _eh_, its
epithelium; _G_, superior muscle of the press; _a_, rachis; _b_, its
posterior enlargement; _c_, infundibulum; _d_, cuticle; _o_, orifice
of the spinning canal; _op_, central canal of the lyre and of the
spinneret; _fi_, hypodermis of the lyre; _f_, _f_, hypodermic pad of
the lyre.
]

The silken substance is then pressed by the more or less powerful
contractions of the muscles of the filator, so that the passage of the
threads is facilitated. If the muscles totally contract, the spinning
canal is opened wide, the threads pass easily upwards and assume the
form of a triangular prism (Fig. 336).

[Illustration:

FIG. 335.—Spinning apparatus, seen from above: _A_, opening of the
spinneret; _B_, central canal of the spinneret (_C_); _D_, common
canal; _E_, canal of Filippi; _F_, excretory canal of a silk-gland;
_i_, orifice of the canal of Filippi’s gland; _l_, rachis; _k_, ring
of the infundibulum; _b_, _c_, _d_, _e_, _f_, cavity of the
different canals; _h_, spur which separates the two excretory
canals.—This and Figs. 331–334 after Blanc.
]

If this contraction diminishes, the chitinous wall of the spinneret
comes together, owing to its elasticity; the ceiling of the canal
approaches the floor; the cavity tends to take the form of a
semicircular slit, and the threads are compressed, flattened. As each
mass or thread of silk is much more voluminous than the canal, except
when the latter is extremely dilated, it follows that the two threads
are always compressed, or squeezed together, and that each of them is
compelled to mould itself in the groove it occupies and to take its
shape. Hence the variations in the appearance of the two masses or
divided portions of silk, which as stated present all grades between the
form of an isosceles-triangular prism and that of a nearly flat ribbon;
but this last case is quite rare. The use of the spinneret, then, is to
compress the thread and to change its form more or less considerably, at
the same time as it diminishes its diameter.

[Illustration:

FIG. 336.—Diagram of the press and its muscles: _a_, lower; _b_,
lateral; _c_, upper muscles of the press.—After Blanc.
]

Moreover, this constant compression of the thread as it passes through
the press keeps it in a certain state of tension so as to allow the
caterpillar while spinning to firmly hold its thread.

Finally, when the worm suspends the contraction of its spinning muscles,
the press flattens, vigorously compresses the thread, and arrests its
motion, in such a way that if there was a strain on the silken fluid
(_bave_), it would break rather than oblige the caterpillar to let go
any more of it.

The press does not act directly on the silken thread, but through the
gummy layer (_grès_) which transmits over the whole surface of the
silken fluid (_brin_) the pressure exerted on it. After having overcome
this difficult passage, the silk thread has acquired its definite form;
it rapidly passes out of the spinneret.

=How the thread is drawn out.=—Having seen, says Blanc, how the two
masses of silk (_brins_), in passing through the spinning apparatus (or
press), join each other, constituting the frothy silken fluid, thus
becoming modified in form, it remains to examine the way in which the
thread is drawn out of the spinneret. If we examine a caterpillar while
spinning, it will be seen that in moving its head it draws on the frothy
mass of silk fixed to the web of the cocoon. This traction certainly
aids very much the exit of the thread, but it is not the only cause.

The silk, Blanc affirms, is pushed out by a force _a tergo_, developed
by different agents, such as the pressure of the distended cuticle or
the silky mass contained in the reservoir, as seen in the section of a
worm which has spun its cocoon. But if we consider a caterpillar before
it has begun to spin, it is difficult to explain the mechanism of
spinning. As Blanc has often observed, in making sections of the heads
of silkworms, two cases arise. Sometimes the worm has already spun a
little, and a certain length of the frothy silk (_bave_) issues from the
orifice of the spinneret, where it forms a small twisted bundle. At
other times the worm has not spun since its last moult or the frothy
mass of silk has broken within the head, and we find the end in the
common tube. In the first of these two cases, the worm, dilating its
press, is able by a general contraction to discharge a little of the
gritty material (_grès_) which lines the ball of silk hanging at the end
of the spinneret. It can also reject a certain quantity of the secretion
of Filippi’s glands and thus soften the gritty substance. The little
plug of silk can then adhere to the body with which it comes in contact.

In the same case it is necessary that the two bits or portions of silk
traverse the press, and this normally has a calibre less than their
diameter. The worm should then distend the spinning tube as much as is
practicable, so as to make the openings as large as possible. It has
been stated that the press is, in this condition, at least as large as
the mass of frothy silk. This Blanc believes (although Gilson thinks
otherwise) is pushed by a force _a tergo_, and reaches the funnel of the
spinning canal; its two bits of silk (_brins_) unite there, penetrate
into the canal itself, and, owing to successive impulses produced by the
general contractions of the worm, press through and pass out of the
spinneret.

While the silkworm is engaged in spinning its cocoon, the spinneret and
press execute very varied movements, determined by the elevator,
depressor, retractor, and protractor muscles of the labium, as well as
those of the press. These movements, originally very numerous, may
combine among themselves, so that the spinneret is susceptible of
assuming during the process of spinning still more diverse positions.

[Illustration:

FIG. 337.—Portion of the silk-gland of _Bombyx mori_: _p_, tunica
propria; _i_, tunica intima; _s_, secretion-cell with branched
nuclei; _a_, separate secretion-cell from the anterior part of the
silk-gland of _Amphidasis betularia_; _b_, the same of _Vanessa
urticæ_; _c_, the same in _Smerinthus tiliæ_.—After Helm.
]

[Illustration:

FIG. 338.—_A_, section of gland of lepidopter: _B_, section of
silk-gland of a saw-fly larva; _n_, nucleus; _i.d_, canals; _d.s_,
cavity.—After Gilson.
]

Histologically the silk-glands are composed of three layers,—the outer,
or _tunica propria_ (Fig 337); the inner, the _tunica intima_; the
middle layer being composed of extraordinarily large epithelial cells
which can be seen with the naked eye, and are also remarkable for the
branched shape of the nuclei (_a_, _b_, _c_, 337), the branches being
more or less lobed, and the larger the cells the more numerous are the
branches of the nucleus. Gilson[53] finds that those of Trichoptera,
Lepidoptera, Diptera, and Hymenoptera ordinarily consist of a small
number of cells; and it is quite common, he says, to find only two cells
in a transverse section (Fig. 338, _A_). In the Tenthredinidæ, however,
“the organ still consists of a tube, the wall of which is composed of
flat cells, but in addition to that, two series of spheroidal cells are
attached to the sides. Each of these cells contains a system of tiny
canals running through their cytoplasm (_B_, _i. d_). These cells are
the secreting elements; they continually cast the silk substance into
the tube.” A peculiarity of the _tunica intima_ is its distinct
transverse striation.

[Illustration:

FIG. 339.—Branching nucleus of spinning gland of Pieris larva.—After
Korschelt, from Wilson.
]

[Illustration:

FIG. 340.—Filippi’s glands (_G_) isolated and seen from above: _e_,
_e_, its lobules; _d_, its excretory canal; _E_, silk-duct; _C_,
common canal; _c_, upper spinning muscle; _b_, lower muscle; _a_,
lateral muscle; _T_, spinneret.—After Blanc.
]

The lining of the glands and of their common duct is moulted when the
caterpillar casts its skin, and this, as well as the mode of
development, shows that the glands are invaginations of the ectoderm.
Gilson finds that the silk-glands and silk-apparatus of Trichoptera are
very similar to those of caterpillars, and that the silk is formed in
the same way.

=Appendages of the silk-gland (Filippi’s glands).=—In most larvæ there
is either a single or a pair of secondary glands which open into the
spinning glands near their anterior end. They are outgrowths of the
gland provided with peculiarly modified excretory cells or evaginations
of the entire glandular epithelium. Those of _Bombyx mori_ (Fig. 340)
are very well developed, and, according to Blanc, form two whitish,
lobulated masses in the labium on each side of the common duct of the
spinning gland. Externally they appear to be acinose; but their
structure, as described by Blanc and by Gilson, is very peculiar. Helm
thinks, with Cornalia, that the function of these glands is to secrete
the adhesive fluid which unites the silk threads, and also to make the
silk more adhesive in the process of spinning, but Blanc states that
this is done before the thread passes into the common excretory tubes,
and he is inclined to think that the secretion serves to lubricate the
spinneret, and thus to facilitate the passage of the thread. On the
other hand, in certain caterpillars these glands are situated quite far
from the spinning apparatus.

The silk-glands in the pupa state undergo a process of degeneration,
and finally completely disappear. They are specific larval organs
evolved in adaptation to the necessity of the insect’s being
protected during its pupal life by a cocoon. (Helm.)

Morphologically the silk-glands are by Lang regarded as modified
coxal glands, and homologues of the setiparous parapodial glands of
chætopod worms, the coxal glands of Peripatus, and the spinning
glands of spiders.

In Scolopendrella, spinning glands are situated in the two last
segments of the body, opening out at the end of the cercopods (Fig.
15, _s.gl_), and the larvæ of the true Neuroptera (Chrysopa,
Myrmeleon, etc.) which spin cocoons, have spinning glands opening
into the rectum. The silk forming the cocoon of the ant-lion, as
Siebold and the older observers have stated, is secreted by the
walls of the rectal or anal sac. Siebold (Anatomy of the
Invertebrates, p. 445) states that in the larva of Myrmeleon, the
silk-apparatus is very remarkable, “for the rectum itself is changed
into a large sac and secretes this substance which escapes through
an articulated spinneret projecting from the opening of the
anus”[54] (Fig. 307, _e_). The larvæ of the Mycetophilidæ have
spinning glands at the hinder end of the body, as also the imago of
the female of the tineid moth Euplocamus. (Kennel.) The larvæ of
ichneumons, wasps, bees, of Cecidomyia, and other Diptera, spin
silken cocoons, but their glands have not yet been examined.

It should also be observed that during the process of pupation the
larvæ of butterflies, of certain flies (Syrphus), and beetles
(Coccinellidæ and some Chrysomelidæ) attach themselves by silk spun
from the anus, so that the pupa is suspended by its tail; such
glands are probably homogenetic with the coxal glands.

The silk in its fluid or soft state is mucilaginous, and according
to Mulder, in the silkworm consists of the following substances,
varying somewhat in their relative proportions by weight:

Silk-fibre material 53.67
Glue (Leim) 20.66
Protoplasm 24.43
Wax 1.39
Coloring matter 0.05
Fat and resin 0.10

LITERATURE ON THE SPINNING GLANDS

=Helm, E.= Anatomische und histiologische Darstellung der
Spinndrüsen der Schmetterlingsraupen. (Zeitschr. f. wissens. Zool.,
xxvi, 1876, pp. 434–469, 2 Taf.)

=Lidth de Jeude, Th. W. van.= Zur Anatomie und Physiologie der
Spinndrüsen der Seidenraupe. (Zool. Anzeiger, 1878, pp. 100–102.)

=Engelmann, W.= Zur Anatomie und Physiologie der Spinndrüsen der
Seidenraupe. (Onderz. Phys. Lab. Utrecht, iii, 1880, pp. 115–119.)

=Joseph, G.= Vorläufige Mitteilung über Innervation und Entwickelung
der Spinnorgane bei Insekten. (Zool. Anzeiger, 1880, pp. 326–328.)

=Poletajew, N.= Ueber die Spinndrüsen der Blattwespen. (Zool.
Anzeiger, 1885, pp. 22–23.)

=Meinert, Fr.= Contribution à l’anatomie des fourmilions. (Overs.
Danske Vidensk. Selsk. Forh. Kjöbenhavn, 1889, pp. 43–66, 2 Pls.)

=Blanc, Louis.= Étude sur la sécrétion de la soie et la structure du
brin et de la bave dans le _Bombyx mori_. Lyon, 1889, pp. 48, 4 Pls.

—— La tête du _Bombyx mori_ à l’état larvaire. Anatomie et
physiologie. (Extrait du volume des Travaux du Laboratoire d’Études
de la Soie. Années 1889–1890, Lyon, 1891, pp. 180, 95 figs.)

=Gilson, G.= Recherches sur les cellules sécrétantes. La soie et les
appareils séricigènes: I. Lépidoptères. (La Cellule, 1890, vi, pp.
115–182, 3 Pls. I, Lépidoptères (suite); II, Trichoptères. Ibid., x,
pp. 71–93, 1893, 1 Pl.)

=Garman, H.= Silk-spinning dipterous larvæ (Science, xx, 1893, p.
215).

Also the writings of Meckel, Pictet, Duméril, Klapálek,
Wistinghausen, Loew, Hagen, Fritz Müller, Kolbe, McLachlan, de
Selys-Longchamps.

_c._ The cæcal appendages.

These diverticula of the mid-intestine (“stomach”) are appended to the
anterior end, and in the living, transparent larva of Sciara, which has
two large, long, slender cœca (Fig. 341), the partly digested food may
be seen oscillating back and forth from the anterior end of the stomach
into and out of the base of each cæcum. In the Locustidæ (Anabrus, Fig.
299) and Gryllidæ (Fig. 344, _e_) there are two large, short cæca, and
in the locusts (Caloptenus) there are six cæca, while cockroaches have
eight. In the Coleoptera (Carabidæ and Dyticidæ) these large cæca appear
to be replaced by very numerous slender, minute villi or tubules, which
arise from the anterior part of the stomach (Figs. 317, _r_, also 342).

These cæca differ in structure from the stomach, as shown by Graber, as
well as by Plateau and by Minot. The latter states that a single
transverse section of one of the diverticula of the locust demonstrates
at once that its structure is entirely different from that of the
stomach.

[Illustration:

FIG. 341.—Larva of Sciara: _s.gl_, salivary gland; _ur.t_, urinary
tubes; _i_ intestine; _st_, stomach; _cae_ cæcal appendages; _t_,
testis.
]

Its inner surface is thrown up into longitudinal folds, generally
twelve in number. These folds shine through the outer walls, and are
accordingly indicated in the drawings of Dufour, Graber, and others.
The entire cæcum has an external muscular envelope, outside of which
are a few isolated longitudinal muscular bands. The folds within are
formed mainly by the high cylindrical epithelium which lines the
whole interior of the cavity. Tracheæ ramify throughout all the
layers outside the epithelium. There are appearances of glandular
follicles in the bottom of the spaces between the folds. (Minot.)

Burmeister supposed that these cæca were analogous to the pancreas, and
this view has been confirmed by Hoppe Seyler, Krukenberg, Plateau, and
others, who claim that the digestive properties of the fluid secreted in
them agrees with the pancreatic fluid of vertebrates.

[Illustration:

FIG. 342—Cross-section of mid-intestine of _Acilius sulcatus_, showing
the arrangement of the cæca, two tracheæ passing into each
cæcum.—After Plateau.
]

_d._ The excretory system (urinary or Malpighian tubes)

The excretory matters or waste products of the blood tissue of worms are
carried out of the body by segmentally arranged tubes called
_nephridia_. As a rule they arise in the blood sinuses of the body and
open externally through minute openings in the skin. As there is a pair
to each segment (in certain oligochete worms two or three pairs to a
segment), they are often called segmental organs. In the annulate worms
each segment of the body, even the cephalic or oral segment, originally
contains a pair of these excretory organs. These vessels may have
survived in myriopods and perhaps do exist in insects as urinary tubes,
and also occur in many of the Arachnida, and thus are characteristic of
each important class of land arthropods, but are either wanting or are
very rudimentary or much modified in the marine classes, notably the
Crustacea and Merostomata (Limulus), where they are represented by the
shell-glands of Copepoda, green glands of the lobster, and the brick-red
glands of Limulus.

[Illustration:

FIG. 343.—Digestive canal of _Perla maxima_: _l_, upper lip; _mh_,
buccal cavity; _ap_, common end of salivary ducts (_ag_); _o_,
œsophagus; _s_, _s_, salivary glands, arranged segmentally; _b_,
cæca of chyle-stomach; _lg_, their ligaments of attachment; _mp_,
urinary tubes; _r_, rectum; _af_, anal orifice.—After Imhof, from
Sharp.
]

In the earliest tracheate arthropod, Peripatus, these tubes are well
developed and are highly characteristic, each segment behind the head
bearing a pair (Fig. 4, _so_{4}_-_so_{9}_). It has been suggested by
some, but not yet proved, that the urinary tubes of insects are
morphologically the same as the segmental organs of worms and of
Peripatus; but there are no facts directly supporting this view, and, as
Sograff states, it is a pure hypothesis and can only be confirmed or
disproved by very detailed researches on the development of the urinary
tubes of myriopods and of insects. Others regard them as probably
homologous with the tracheæ, since they have a similar origin. As,
however, they arise in the embryo as outgrowths of the proctodæum they
may have arisen in myriopods and insects independently, and not be
vermian heirlooms.

While in worms and in Peripatus a pair of these segmental organs occur
in each segment, in insects this serial arrangement is not apparent;
those with a purely excretory function are not segmentally arranged,
with outlets opening externally, but arise as outgrowths of the
hind-intestine or proctodæum of the embryo, not being segmentally
arranged. The place of their origin is usually the dividing line between
the mid and hind intestine (Fig. 343, _mp_); this applies to
Scolopendrella (Fig. 15, _urt_) as well as to insects.

The urinary tubes are usually long, slender, blind, tubular glands
varying in number from two to over a hundred, which generally arise at
the constriction between the mid and hind intestine, and which lie
loosely in the cavity of the body, often extending towards the head, and
then ending near the rectum (Figs. 301, 310, _vm_). They were first
discovered by the Italian anatomist Malpighi, after whom they were
called the Malpighian tubes. While at first generally regarded as
“biliary” tubes, they are now universally considered to be exclusively
excretory organs, corresponding to the kidneys of the higher animals.

[Illustration:

FIG. 344.—Digestive canal and appendages of the mole-cricket; _a_,
head: _b_, salivary glands and receptacle; _c_, lateral pouch; _d_,
stomatogastric nerves; _e_, anterior lobes of stomach; _f_, peculiar
organ; _g_, neck of stomach; _h_, plicate part of same; _i_, rectum;
_k_, anal gland; _m_, urinary tubes.—After Dufour, from Sharp.
]

Usually arising from the anterior end of the hind-intestine where it
passes into the mid-intestine, in certain forms they shift their
position, in some Hemiptera (Lygæus, Cimex) opening into the rectum,
while in the Psyllidæ they arise from the slender hinder part of the
mid-intestine, being widely separated at their origin. (Fig. 321.)

The length varies in different groups; where they are few in number (two
to four, six to eight), they are very long, but where very numerous they
are often short, forming dense tufts, each tuft connecting with the
intestine by a common duct (ureter), or, as in the mole-cricket, the
numerous tubes empty into a single duct (Fig. 344); in the locusts
(Acrydiidæ), however, they are arranged in 10 groups, each group
consisting of about 15 tubes, making about 150 in all; and are much
convoluted and wound irregularly around the digestive canal, and when
stretched out being about as long as the entire body.

The urinary tubes occur in twos, or in multiples of two, though a
remarkable exception is presented in the dipterous genera Culex and
Psychodes, in which there are five tubes; the young and fully grown
larvæ, as well as the pupa and imago of Culex, having this number (Fig.
433, _mg_.)

In many insects (Pentatoma, Cimex, Velia, Gerris, Haltica, Donacia,
and often in caterpillars), the vessels open into a sort of urinary
bladder connecting with the intestine on one side.

[Illustration:

FIG. 345.—_A_, section of urinary tube of Periplaneta; _B_, part of
tube of Perla; _p_, peritoneal membrane; _c_, cavity or lumen;
_n_, nucleus of a secreting cell.—After Schindler.
]

In the larvæ of some insects the blind ends of the tubes are often
externally bound to the rectum, in the silkworms being attached by
fine threads to the intestine, while in some flies (Tipula and
Ctenophora), two vessels may unite to form a loop. In all larval
Cecidomyiæ, the two tubes are united to form a loop which curves
backward, opening near the vent, the proctodæum being very short.
(Giard.)

[Illustration:

FIG. 346.—Portion of a urinary tube of _Calliphora vomitoria_: _tr_,
trachea; _l_, lumen; _k_, nucleus.—After Gegenbaur.
]

While usually the urinary vessels form simple tubes, in many species
of Lepidoptera and Diptera they are branched, thus resembling those
of spiders and scorpions. Moreover, in many Lepidoptera and Diptera
(Fig. 308), the tubes are not simple, but are lobulated, and in some
Hemiptera (Pentatoma, Notonecta, and Tettigonia) are twisted or
lace-like. In rare cases there are two kinds of urinary tubes; in
_Melolontha vulgaris_, two of them are partly lobulated and yellow,
while the other two are simple and white. Their color in beetles
varies, some being whitish or yellowish; in Geotrupes, Dyticidæ,
Hydrophilidæ, etc., reddish brown; in Gryllotalpa as well as
_Locusta viridissima_, there are two different kinds of vessels,
differing in contents and in color (white or yellow), as well as
histologically. (Schindler.)

The exterior of the tubes is richly provided with tracheæ, which
often form a web around them, and the fine branches often seem to
attach them to the intestine. In Acheta they are enveloped by a very
delicate, loose network of muscular fibres. (Schindler.)

The urinary tubes consist, according to Schindler, of at least three
cellular layers (Fig. 345):—

1. An external, connective, nucleated membrane, the peritoneal
membrane.

2. A very delicate homogeneous basal membrane, the _tunica propria_.

3. A single layer of large polygonal excretory cells.

4. Lining the internal canal a chitinous layer penetrated by
pore-canals, the _intima_ often wanting.

The secretory cells are usually of the same size, but in many cases
are relatively small; sometimes four to six or more form the
periphery of the canal, sometimes three or only two. In some insects
the cells are so very large that a single cell forms the entire
periphery. The nuclei in the Lepidoptera (Papilio, Pontia, Cossus)
are large and irregularly branched.

The excretions of the Malpighian vessels, derived from the blood and
from the fat-body, are more or less fluid and granular, sometimes
pulpy. From the cells they pass into the canal, thence into the
intestine, and thence out of the body. How, says Kolbe, the
secretion passes into the intestine, whether by the contraction of
the fine fibrillæ of the peritoneal membrane, or by the external
pressure of the other organs, or by the pressure of the secretory
matter behind, is not yet known. Grandis observed in living
Hydrophilus that the urinary tubes moved, without the muscles
seeming to show what caused the motion. Moreover, the cells
incessantly changed their form. At a lower temperature such motions
ceased. The tracheæ, ending freely in the cells, did not anastomose.
(Kolbe.)

The different colors of the tubes (white, yellow, red, brown, or
green) is due to the hue of the excretions, and is independent of
the color of the blood and of the urinary substances held in the
secreted matter.

Schindler found that insects of different stages, collected in
winter, differed very much in their urinary secretions, the tubes in
the adults being entirely empty, while in the larvæ they were filled
full, so that he concluded that in the former the process of
excretion during the winter hibernation is very slow, but in the
latter very rapid.

As to the activity of the urinary vessels the following experiments
will throw some light. Tursini fed a Pimelia with fuchsin; its
urinary tubes were consequently colored red. Schindler fed insects
with indigo-carmine, which was excreted by the urinary tubes;
Kowalevsky arrived at the same results, which seems to prove that
these vessels are analogous to the kidneys of vertebrates. Moreover,
Schindler injected through the side of the first abdominal segment
into the cavity of the body of a Gryllotalpa a concentrated solution
of sodium salt of indigotin-disulphonic acid. After one or two hours
the external portion of the epithelium of the urinary vessels was
stained deep blue, while the inner portion remained of the normal
transparency; the nuclei being for the most part deeply stained.
Between one and two days after, the staining matter had not yet
wholly passed through the central canal, the surface recently
stained still appearing light blue.

The solid contents of the urinary tubes consist partly of crystals,
which occur singly in the epithelial cells, or form scattered masses
when situated in the central canal. Besides tabular rhombic crystals,
there occur concretions which contain uric acid, and probably consist of
urate of soda, also octahedral crystals of chloride of soda, and
quadro-pyramidal crystals of oxalate of lime. Also acicular prisms
occur; besides chloride of soda, phosphates, carbonate of lime, oxalate
of lime in quantity, leucine, coloring matters, etc.; while the fluid
secretion also contains urea (?), uric acid, and abundant urates; uric
acid crystals were precipitated by the addition of acetic acid, and by
adding hydrochloric acid crystals belonging to the dimetric system were
formed. The often numerous spheroidal small granules are biurate of soda
and biurate of ammonia. Pale, concentrically banded concretions are
leucine pellets.

According to Kölliker the contents of the urinary vessels[55] in
general are: (1) round granules of urate of soda and urate of
ammonia; (2) oxalate of lime; and (3) pale transparent concretions
of leucine. Crystals of taurin are also said to occur. (Claus’
Zoölogy, p. 531.)

Although uric acid is characteristic of the urinary tubes, yet
sometimes it is wanting in them, while uric acid substances in
quantity occur in the fat-body or in the mid-intestine.

In the living insect the urinary tubes remove urates from the blood;
“the salts are condensed and crystallized in the epithelial cells,
by whose dehiscence they pass into the central canals of the tubules
and thence into the intestine.” (Miall and Denny.)

The process of excretion is carried on not only by the urinary
tubes, but also, as Cuénot has recently shown (1896) in Orthoptera,
by the pericardial cells and certain cells of the fat-bodies. In the
last-named cells urates are stored throughout life; the pericardial
cells apparently secrete but do not store waste products, which are
finally eliminated by the urinary tubes, the latter constantly
eliminating waste.

=Primitive number of tubes.=—Wheeler considers the primitive number
of urinary tubules to be six, other authors regarding two pairs as
the primary or typical number; and while Wheeler agrees that the
more ancestral tracheate arthropods had but a single pair,
Cholodkowsky supposes the primitive number in insects themselves to
be a single pair. This view is strengthened by the fact that
Scolopendrella has but a single pair (Fig. 15).

While Peripatus has no urinary tubes, in Myriopods a single pair
arises, as in insects, from the hind-intestine.

[Illustration:

FIG. 347.—Section of proctodæum of embryo locust, showing origin of
urinary tubes (_ur.t_); _ep_, epithelial or glandular layer; _m_,
cells of outer or muscular layer; _a_, section of a tube.
]

When in insects the number of these tubes is few, they are, with
rare exceptions, arranged in pairs, so that Gegenbaur and others
have considered this paired arrangement as the primitive one. When
the tubules are very numerous in the adult, as in Orthoptera, the
embryos and larvæ have a much smaller number, Wheeler stating that
“in no insect embryo have more than three pairs of these vessels
been found.” We have observed 10 primary tubes in the embryo of
Melanopus (Fig. 347), from each of which afterwards arise 15
secondary tubules. In the Termites, only, do the young forms have
more urinary tubes than the adults.

In Campodea there are about 16 urinary tubes and in Machilis either
12 (Grassi) or 20 (Oudemans); but in other Thysanura the number is
much less, Lepisma having either four, six, or eight, according to
different authors, and both Nicoletia and Lepismina having six,
opening separately into the hind-intestine. On the other hand, these
organs have not yet been detected in Japyx. Whether they exist at
all in the Collembola, which are degenerate forms, is doubtful. The
weight of opinion denies their existence, though they may yet be
found existing in a vestigial condition. They are said by Tullberg
and by Sommer to exist in Podura, but are of peculiar shape.

Coming now to the winged insects, in what on the whole is perhaps
the lowest or most generalized order, the Dermaptera, the number is
over 30, and their insertions regularly encircle the intestine.
(Schindler.) In the most ancient and generalized family of
Orthoptera, the Blattidæ, Schindler detected from 60 to 70 tubes,
but in a nymph of Periplaneta not quite 10 mm. in length he found
from 16 to 18, and in nymphs 4 to 5 mm. long there were only eight
vessels; while Wheeler has found in the embryo of _Phyllodromia
germanica_ but four tubes. In the adult Acrydiidæ there are as many
as 150, in the Locustidæ between 40 and 50, and in the Gryllidæ
about 100.

The Ephemeridæ with about 40, the Odonata with 50 to 60 tubules, the
Perlidæ with from 50 to 60, are polynephrious; while the Termitidæ
and Psocidæ are oligonephrious, the former having from six to eight
and the Psocidæ only four tubes. So also all the other orders not
mentioned, except the Hymenoptera, have few of these tubes. The
Hemiptera, with none in Aphidæ, a single pair in the Coccidæ, and
two in all the rest of the order, have the fewest number.

In the Neuroptera there are from six to eight, while in a larva,
possibly that of Chauliodes, Wheeler finds the exceptional number of
seven.

The closely allied order Mecoptera (Panorpidæ), and also the
Trichoptera, agree with the Neuroptera (Sialis) in having six.
According to Cholodkowsky all Lepidoptera have six of these vessels,
except Galleria, which has but four. He finds that in _Tinea
biselliella_ (also _T. pellionella_ and _Blabophanes rusticella_)
the larva has six vessels, which, however, undergo histolysis during
pupation, a single pair arising in their stead. On this account he
regards the primitive number of urinary tubes as two, or a single
pair, this return from six vessels in the larva to two in the imago
being considered a case of atavism.

In the Coleoptera, the number of urinary tubes is from four to six;
in what few embryo beetles have been examined (Doryphora,
Melolontha), there are six vessels, but in the embryo of _Dyticus
fasciventris_, Wheeler has detected only four, this number being
retained in the adult. He thinks that in beetles in general, a pair
of vessels must be “suppressed during post-embryonic development,
presumably in early larval life.”

In Diptera and Siphonaptera, the number four is very constant, there
being, however, a fifth one in Culex and Psychoda (Fig. 400.)

The number of these vessels is very inconstant in the Hymenoptera,
varying from six (Tomognathus, an ant, worker) to 12 (Myrmica), and
in Apis reaching the number of 150.

In the embryo of the honey-bee and wall-bee (Chalicodoma), there are
only four; we still lack any knowledge of the number in embryo
saw-flies.

The following is a tabular view of insects with few urinary tubes
(Oligonephria) and many (Polynephria). It will be seen that the number
has little relation to the classification or phylogeny, insects so
distantly related as the Orthoptera and Hymenoptera being
polynephrious:—

_Oligonephria_

Collembola, 2 (Podura), Tullberg and also Sommer.
Thysanura, 4 (Lepisma); in Campodea, 16; in Machilis, 12 or 20; wanting
in Japyx.
Psocidæ, 4.
Termitidæ, 6 (many in the young, Rathke).
Mallophaga, 4.
Physapoda, 4.
Hemiptera, 2 (Coccidæ, none in Aphidæ).
Neuroptera, 6–8. (In Sialidæ and Rhaphididæ 6; in Myrmeleonidæ and
Hemerobiidæ, 8).
Trichoptera, 6.
Mecoptera, 6.
Lepidoptera, 2–4–6 (2 in Tinea, Tineola, and Blabophanes; in Pterophorus
and Yponomeuta, 4).

_Coleoptera_, 4–6; never more.

4
Carabidæ,
Dyticidæ,
Staphylinidæ,
Gyrinidæ,
Palpicornes,
Lamellicornes,
Cantharidæ,
Buprestidæ
(in larva, 6; in beetle, 4).

6
Byrrhidæ,
Nitidulidæ,
Dermestidæ,
Cleridæ,
Meloidæ,
Pyrochroidæ,
Bruchidæ,
Bostricidæ,
Cerambycidæ
Chrysomelidæ,
Coccinellidæ.

Diptera, branching into 4 (Gegenbaur); in Culicidæ, and Psychoda, 5.
Siphonaptera, 4.

_Polynephria_

Orthoptera, 100–150. (In embryo Blattids, 4; in embryo locust, 10; in
nymph of Gryllotalpa, 4.)
Dermaptera, “over 30” (Schindler).
Perlidae, 50–60.
Plectoptera (Ephemeridæ), 40.
Odonata, 50–60.
Hymenoptera, 20–150. (In embryo bees only 4; Cynipidæ, Ichnenumonidæ,
and Formicidæ have the smallest number, 6–12.)

Here should be mentioned the singular fact discovered by Koulaguine
that in the larva of Microgaster, the urinary tubes have no
connection with the intestine, but open dorsally on the outside of
the body on each side of the anus. Ratzeburg had stated that the
last segment of the body was in the form of a vesicle. Koulaguine
now shows that this vesicle is in reality the end of the intestine
opening upwards; as the result of this dorsal opening of the
intestine the Malpighian vessels open on the sides of the oval vent,
and have no connection with the intestinal canal. Whether this is of
morphological import, or is only a secondary adaptation, Koulaguine
does not state, his paper being a preliminary abstract.

Wheeler thus sums up our present knowledge regarding the number and
homologies of the Malpighian or urinary tubes:

1. It is very probable that the so-called Malpighian vessels of
Crustacea and Arachnida are not the homologues of the _vasa
Malpighi_ of the Eutracheata (insects and myriopods).

2. The Malpighian vessels of the Eutracheata arise as paired
diverticula of the hind-gut and are, therefore, ectodermal.

3. In no insect embryo are more than six vessels known to occur;
although frequently only four are developed.

4. The number six occurs either during embryonic or post-embryonic
life in members of the following groups: Apterygota, Orthoptera,
Corrodentia; Neuroptera, Panorpata, Trichoptera, Coleoptera,
Lepidoptera, and Hymenoptera.

5. The number four seems to be typical for the Corrodentia,
Thysanoptera, Aphaniptera, Rhynchota, Diptera, and Hymenoptera.

6. The embryonic number in Dermaptera, Ephemeridea, Plecoptera, and
Odonata has not been ascertained, but will probably be found to be
either four or six.

7. There is evidence that in at least one case (Melolontha), the
tetranephric is ontogenetically derived from the hexanephric
condition by the suppression of one pair of tubules.

8. It is probable that the insects which never develop more than
four Malpighian vessels have lost a pair during their phylogeny.

9. The post-embryonic increase in the number of Malpighian vessels
in some orders (Orthoptera, Odonata, Hymenoptera) is secondary and
has apparently arisen to supply a demand for greater excreting
surface.[56]

LITERATURE ON THE EXCRETORY (URINARY) ORGANS

=Malpighi, M.= Dissertatio epistolica de Bombyce, Societati regiæ
Londini ad scientiam naturalem promovendam institutæ dicata.
(Londini, 1669, 12 Pls.)

=Herold, M. J. D.= Entwicklungeschichte der Schmetterlinge. 1815.

=Rengger, J. R.= Physiologische Untersuchungen über den tierischen
Haushalt der Insekten. Tübingen, 1817, pp. 82.

=Wurzer.= Chemische Untersuchungen des Stoffes in den Gallgefässen von
_Bombyx mori_. (Meckel’s Archiv f. Physiol., iv, 1818, pp. 213–215.)

=Gaede, H. M.= Physiologische Bemerkungen über die sogenannten
Gallgefässe der Insekten. (Nova Acta Acad. Caes. Leopold.-Carolin.,
1821, x, Pars II, pp. 186–196.)

=Meckel, J. F.= Ueber die Gallen- und Harnorgane der Insekten.
(Meckel’s Archiv, i, 1826, pp. 21–36.)

=Audouin, J. V.= Calculs trouvés dans les canaux biliaires d’un cerf
volant. (Ann. sc. nat., 2 Sér., 1836, v, pp. 129–137.)

=Frey und Leuckart.= Anatomie und Physiologie der Wirbellosen. 1843.

=Dufour, L.= Mémoire sur les vaisseaux biliaires ou le foie des
Insectes. (Ann. sc. nat., 1848, Sér. 2, xix, pp. 145–182, 4 Pls.)

=Karsten, H.= Harnorgane von _Brachinus complanatus_. (Müller’s Archiv
f. Anat. und Physiol., 1848, pp. 367–374.)

=Fabre, J. L.= Étude sur l’instinct et les metamorphoses des
Sphégiens. (Ann. d. sc. nat., 4 Sér., 1856, vi, pp. 137–189.)

—— Étude sur le rôle du tissu adipeux dans la sécrétion urinaire chez
les Insectes. (Ibid., 4 Sér., xix, pp. 351–382.)

=Schlossberger, J. E.= Untersuchungen über das chemische Verhalten der
Krystalle in den Malpighischen Gefässen der Raupen. (Archiv f. Anat.
und Physiol., 1857, pp. 61–62.)

=Leydig, F.= Lehrbuch der Histiologie. 1857.

=Sirodot, S.= Recherches sur les sécrétions chez les Insectes. (Ann.
sc. nat., 4 Sér., Zool., 1858, x, pp. 141–189, 251–334, 12 Pls.)

=Kölliker, A.= Zur feineren Anatomie der Insekten (Ueber die
Harnorgane, u.s.w.) (Verhandl. d. Physikal.-medizin. Gesellsch. in
Würzburg, viii, 1858, pp. 225–235.)

=Schindler, E.= Beitrage zur Kenntnis der Malpighischen Gefässe der
Insekten. 3 Taf. (Zeitschr. f. wiss. Zool., xxx, 1878, pp. 587–660.)

=Chatin, G.= Note sur la structure du noyau dans les cellules
marginales des tubes de Malpighi chez les Insectes et les
Myriapodes. (Ann. d. sc. nat., 6 Sér., xiv., 1882, pp. 7, 1 Pl.)

=Witlaczil, E.= Zur Anatomie der Aphiden. (Arbeiten a. d. Zool.
Instit. d. Univers. Wien., iv, 1882, pp. 397–441, 3 Taf.)

=Cholodkowsky, N.= Sur les vaisseaux de Malpighi chez les
Lépidoptères. (Compt. rend. Acad. d. Sc., Paris, xcix, 1884, pp.
631–633.)

—— Sur la morphologie de l’appareil urinaire des Lépidoptères.
(Archives de Biologie, 1887, vi, pp. 497–514, 1 Pl.)

=Loman, J. C. C.= Ueber die morphologische Bedeutung der sogenannten
Malpighischen Gefässe der echten Spinnen. (Tijdschr. Nederl. Dierk.
Ver. (2) Deel 1, 1887, pp. 109–113, 4 Fig.)

=Marchal, P.= Contribution à l’étude de la désassimilation de l’azote.
L’acide urique et la fonction rénale chez les Invertébrés. (Mém.
Soc. Zool. de France, 1889, iii, pp. 42–57.)

=Kowalevsky, A. O.= Ein Beitrag zur Kenntnis der Exkretionsorgane.
(Biol. Centralbl., ix, 1889–90, pp. 33–47, 65–76, 127–128.)

—— Sur les organes excréteurs chez les arthropodes terrestres.
(Congrès international de Zool., 2^{me} Session à Moscou, 1892, Pt.
I, pp. 186–235, 4 Pls.)

=Griffiths, A. B.= On the Malpighian tubules of _Libellula depressa_.
(Proc. Roy. Soc., Edinburgh, 1889, xv, pp. 401–403, Figs.)

=Grandis, V.= Sulle modificazioni degli epitelii ghiandolari durante
la secrezione. (Atti Accad. Torino, 1890, xxv, pp. 765–789, 1 Pl.;
Archiv Ital. Biol., 1890, xiv, pp. 160–182, 1 Pl.)

=Koulaguine, N.= Notice pour servire à l’histoire du développement des
hyménoptères parasites. (Congrès internat. de Zool., 2^{me} Session
à Moscou, 1892, Pt. I, pp. 253–277.)

=Sograff, Nicolas.= Note sur l’origine et les parentés des
Arthropodes, principalement des Arthropodes trachéates. (Congrès
internat. de Zool., 2^{me} Session à Moscou, 1892, Pt. I, pp.
278–302.)

=Giard, Alfred.= (Note on the urinary tubes of larval Cecidomyia.
Annals Ent. Soc., France, lxii, 1893, pp. lxxx-lxxxiv, 1 Fig.)

=Wheeler, William M.= The primitive number of Malpighian vessels in
insects. (Psyche, vi, May-December, 1893, Parts 1–6, pp. 457–460,
485–486, 497–498, 509–510, 539–541, 545–547, 561–564.)

=Metalnikoff, C. K.= Organes excréteurs des insectes. (Bull. Acad.
imp. Sci. St. Pétersbourg, 1896, iv, pp. 57–72, in Russian, 1 Pl.)

See also the works of Straus-Dürckheim, Will (Müller’s Archiv. 1848,
p. 502), Brugnatelli, Leidy, Dufour, Ramdohr, Basch, Davy, Grassi,
Minot, Berlese, Adlerz, Marchal (Bull. Ent. Soc. France, 1896, p.
257); Bordas (Appareil glandulaire des Hyménoptères, 1894), also C.
R. Acad. Sc. Paris, 1897.

_e._ Poison-glands

Poison-glands are mainly confined to the stinging Hymenoptera, _i.e._
certain ants, and the wasps and bees, but also occur in the mosquito,
while many, if not most bugs, seem to instil a drop of poison into the
punctured wounds they make.

In the honey and other bees the poison apparatus consists of two
poison-glands whose secretion passes by a single more or less convoluted
efferential duct into the large poison-sac, and thence by the excretory
duct, which is enlarged at the base of the sting (Figs. 194, 195), out
through the sting by the same passage as the eggs. According to Carlet,
the poison apparatus of bees consists of two kinds of glandular organs,
of which one kind secretes a feebly alkaline fluid, the other an acid
product. The poison is only effective when both fluids are mixed. The
resultant venom is always acid. The action of this venom upon some
animals, as rabbits, frogs, and certain beetles, is slight; but the
domestic fly and the flesh-fly are immediately killed by it. The
inoculation of a fly with the secretion of one of the glands does not
produce death until after a considerable time, but death follows very
quickly if the same fly is subjected to a second inoculation, this time
with the secretion of the other gland. The alkaline glands are in bees
and all poisonous Hymenoptera strongly developed, but become vestigial
in those forms which sting their prey to serve as food for their larvæ.
The poison which the solitary sand and wood wasps and Pompilidæ inject
into their victims only paralyzes them.

[Illustration:

FIG. 348.—The poison apparatus of Ichneumon: _T_, sting; _GA_ acid
gland; _TG_, _R′_, its tubes opening into the common poison-sac or
reservoir; _ce_, its efferent canal; _Ga_, the tubular alkaline
gland; _R_, the glandular end; _a_, the reservoir; _ce_, its duct;
_Gac_, the accessory gland—After Bordas.
]

[Illustration:

FIG. 349.—Cephalic gland of Belostoma.
]

Bordas has found both the alkaline gland (gland of Dufour) and the
acid gland to occur in a hundred species of Hymenoptera, including
not only Aculeata, but also Ichneumonidæ (Fig. 348), Tenthredinidæ,
and they may be safely said to be of general occurrence. The acid
gland consists of three parts, the glandular portion, the reservoir
for the poison, and the secretory canal. The alkaline gland is an
irregular tube, with a striated surface and without a reservoir. In
most Hymenoptera there is still a third gland, which is unpaired,
granular, rectangular or lanceolate, with a short filamentous duct
which opens beside the orifice of the alkaline glands.

The poison in ants, wasps, and bees consists of two substances, _i.e._
formic acid and a whitish, fatty, bitter residue in the secretion of the
glands; the corroding active formic acid is the essential part of the
poison. (Will.)

In Melipona the sting and poison-glands are aborted; in certain ants
(Formica, Lasius, etc.) the sting is wanting, but the poison-sac is
extraordinarily large.

Bordas finds in various species of Ichneumon three kinds of glands
opening into the base of the sting. The first two correspond to the
acid (Fig. 348, _G.A_) and alkaline (_G.A_) glands of bees and wasps
(Vespidæ, etc.), and the third (_G.ac_) is situated between the two
lateral muscular bundles which attach the base of the sting to the
last abdominal segment. The poison-reservoir (Fig. 348, _V_) is
recognized by its yellow color and diaphanous and striated
appearance. It is situated on the left of the hind-intestine, a
little in front of the rectum. The tubular gland (_Ga_) or alkaline
gland of aculeate Hymenoptera is remarkably large; it is situated on
the left side of the body. The accessory gland (_G.A_) is elongated,
triangular, flat, its duct opening at the base of the alkaline
gland; it is formed of small spherical cells. Bordas has met with
well-developed poison-glands in forty species belonging to the
Terebrantia, including that of Tenthredo, Emphytus, as well as
various genera of Ichneumonidæ, but in all these species the
accessory gland was wanting.

[Illustration:

FIG. 350.—View from above of the cephalic gland of Belostoma, ×
20.—This and Fig. 349 after Locy.
]

Under the name of cephalic glands (Fig. 349), Locy describes a pair of
glands in the head of Nepidæ. The epithelial or secreting cells are
8–sided (Fig. 350). “When these insects are irritated,” he says, “a
secretion is freely thrown out around the base of the beak, which
produces death very quickly when introduced on a needle point into the
body of an insect.” He infers that the cephalic glands may be the source
of this poisonous secretion. The poisonous salivary fluid of the larva
of Dyticus is referred to on p. 324.

That the mosquito injects poison into the wound it makes has been proved
by Macloskie, who discovering fine droplets of a yellow oily-looking
fluid escaping from the end of the hypopharynx, afterwards detected the
poison-glands. It appears that the two salivary glands are subdivided,
each into three lobes, the middle of which (Fig. 351, _pg_) differs from
the others in having evenly granulated contents and staining more deeply
than the others. Having examined the preparations, we agree with the
discoverer that these lobes secrete the poison. The poison is diluted by
the secretion of the salivary lobes, and the two efferent ducts, one
from each set of glands, “carry forward and commingle the
venomo-salivary products in the main duct; and the stream is then
carried by the main duct to the reservoir at the base of the
hypopharynx.”

[Illustration:

FIG. 351.—_A_, median section of head, showing (_du_) the
venomo-salivary duct, with its insertion in (_hy_) the hypopharynx;
_cb_, brain; below is the pharyngeal pump, leading from (_œ_) the
œsophagus; _lre_, base of labrum-epipharynx; _m_, muscle; _n_,
commissure (other parts removed). _B_, the venomo-salivary duct,
showing its bifurcation, and the three glands on one of its
branches; _pg_, poison gland; _sg_, the upper of the two salivary
glands. _C_, the bifurcation of the duct, with its nucleated
hypodermis.—After Macloskie.
]

_f._ Adhesive or cement-glands

Dewitz has discovered in ants and bees, in close connection with the
poison-glands, and like them discharging their secretion through the
sting, cement-glands. They arise by budding at the base of the
poison-glands.

The two glands in these Hymenoptera correspond to the tubular glands of
the Orthoptera, which open at the base of the inner sheath of the
ovipositor (Fig. 299, _sb_), so that the secretion flows out through it
as the poison of bees, etc., out of the sting. The use of the secretion
of these glands is either to glue the eggs together, or to afford
material for the egg-case of cockroaches and Mantidæ and the gummy
egg-case of the locusts, etc. The contents of the cement-glands serves
for the fixture of the eggs after deposition. In the stinging
Hymenoptera one of the cement-glands is an accessory gland; the other
becomes the poison-sac. The cement-glands are in the Hemiptera only
short blind sacs, in the Lepidoptera and Diptera long convoluted tubes,
tubular and branched in the Coleoptera, or richly branched in the
Ichneumonidæ and Tenthredinidæ. In the cockroach there are two
cement-glands, but the right one is probably of no functional
importance. The left one is filled with a milky substance, containing
many crystals and a coagulable fluid, out of which the egg-capsule
(oötheca) is formed. (Miall and Denny.) In the locusts the sebific or
cement-gland (Fig. 298, _sb_) secretes a copious supply of a sticky
fluid, which is poured out as the eggs pass out of the oviduct and
agglutinates the eggs into a mass, forming a thin coating around each
egg, which from the mutual pressure of the eggs causes the tough coating
to be pitted hexagonally. In other insects also (Trichoptera,
Chrysopidæ, Lepidoptera, etc.) there are similar secretions for the
protection or fastening of the eggs when laid.[57] The Trichoptera lay
their eggs either in or on the surface of the water in bunches or in
strings or in annular gelatinous masses on stones or on plants. This
jelly-like substance is secreted by two highly developed paired anal
glands. (Weltner, in Kolbe, p. 621.) Also in certain dragon-flies
(Libellula, Diplax, and Epitheca) the eggs are laid in jelly-like
masses.

With a similar secretion, spun from the end of the abdomen, the Psocidæ
cover their little bunches of eggs laid on the under side of leaves; and
the silk thread forming the egg-sac of the great water-beetle
(Hydrophilus) is secreted from such anal glands.

_g._ The wax-glands

Besides the honey-bee, which secretes wax in little scales on the under
side of the abdomen, the bodies of many other insects, such as the plant
and bark lice, as well as the Psyllidæ, Cicadidæ (especially Flata and
Lystra), are covered with a waxy powder, or as in Chermes, Schizoneura,
Flata, etc., with wool-like filaments of wax.

[Illustration:

FIG. 352.—Under side of worker honey-bee, carrying wax scales, ×
3.—After Cheshire.
]

[Illustration:

FIG. 353.—Nymph of Lachnus, showing position of wax-glands.—Gissler
_del._
]

The wax is secreted by minute unicellular dermal glands, which in the
lower insects (Hemiptera) are distributed nearly all over the body, but
in the bees are restricted either to the under (Apis, Fig. 352) or upper
side (Trigona) of the end of the abdomen.

The wax-glands of Pemphigus, Chermes, etc., lie under the little warts,
seen in _Lachnus strobi_, the white-pine aphis, to be distributed in
transverse lines across the back and sides of the abdominal segments
(Fig. 353). These warts are surrounded by a chitinous ring, and divided
into delicately marked areas. Through the delicate numerous pits in the
chitinous membrane of these areas the little waxen threads project,
since under each area ends a duct leading from a large glandular cell,
which is a specially modified hypodermis cell (Claus). The wax threads
are hollow, and all those arising from a single glued cell form a
bundle, whose threads separate from each other and form a white woolly
down or bloom covering the body. Witlaczil also shows that gall-forming
Aphids secrete a wax-like substance, which, during the movements of the
insects in the gall, is rubbed off, becoming a watery layer mixed with
the fluid excrement, which forms a spherical impervious layer lining the
gall, and thus rendering possible the mode of life of the gall-lice.

In the Psyllidæ Witlaczil has discovered wax-glands which also secrete
slender waxen threads. They are situated in groups of two or three at
the end of the abdomen near the anus, and arise from hypodermis cells.
The wax threads surround the liquid excrement as it passes out of the
vent, covering it with a continuous layer of wax. The excrement
accordingly is discharged very slowly and gradually, in sausage-shaped
masses slightly strung together and rolled into close spirals. The body
becomes unavoidably smeared with the sticky excrement, since it is not
entirely covered by the waxy layer. Moreover, in the larvæ of many
Psyllidæ waxen threads are formed on the upper side of the abdomen; they
are for the most part tightly curled or frizzly, like wool, and form,
though partly torn, a waxen coat, chiefly on the side and back of the
thorax and abdomen. The insects appear therefore as if covered with
dust. The mature animals of many species are also covered with a waxen
down. The wax threads rapidly dissolve and disappear in alcohol. From a
wax-like substance more or less easily dissolved in alcohol arise
peculiar hair-like structures which, in the larvæ of Psyllidæ, are
situated on the side and end of the body and also on the rudiments of
the wings. They are readily distinguished from ordinary hairs, as they
arise from glandular cells, and are of very different lengths, more or
less like bristles, but hollow, and very brittle. They are leaf-like in
the first nymphal stages of _Trioza rhamni_, but in following stages
become narrow and form a row around the entire periphery of the body.

The waxen dorsal shield which protects the body of bark-lice (Coccidæ)
is a similar product.

[Illustration:

FIG. 354.—Young nymph and developing scale of _Aspidiotus
perniciosus_: _a_, ventral view of nymph, showing sucking beak with
setæ separated, with enlarged tarsal claw at right; _b_, dorsal view
of same, somewhat contracted, with the first waxy filaments
appearing; _c_, dorsal and lateral views of same, still more
contracted, illustrating further development of wax secretion; _d_,
later stage of same, dorsal and lateral views, showing matting of
wax secretions and first form of young scale; all greatly
enlarged.—After Howard and Marlatt, Bull. 3, N. S., Div. Ent., U. S.
Dept. of Agr.
]

Witlaczil has described the way it is formed in Aspidiotus and
Leucaspis. The freshly hatched nymph shows no signs of a waxy
secretion. But eventually waxen threads arise first on the hinder
and anterior end of the body, and then over the whole surface. These
threads interlace into a sort of felting and thus form the shield,
which is usually much larger than the body and lies closely upon it.
The shield is formed after the first moult. It is noteworthy that
these threads are matted together to form as thick a tissue as that
of the shield itself. The shield is whitish or gray and rather thin.
On the thinnest part of the edge the single threads may be drawn
out. The growth of the shield advances with the increase in size of
the nymph around the entire edge, but is greatest behind. The first
two larval skins are retained on the back under the shield. Also a
very thin waxen pellicle remains on the resting place of the insect
when it is raised. The wax-glands open in the pitted fields, and
appear as clear brownish cells which are distinguished from the
ordinary hypodermis cells by their greater size. (Witlaczil. Compare
also Fig. 354.)

[Illustration:

FIG. 355.—Wax disks of social bees: _a_, _Apis mellifica_, worker;
_b_, do., queen; _c_, Melipona, worker; _d_, Bombus, worker.—From
Insect Life, U. S. Dept. Agr.
]

The wax-glands in the honey-bee are scale-shaped organs situated on the
under side of the four last abdominal segments (Fig. 355). These secrete
the wax, which appears as whitish scales, and secretion is only possible
when the bees have sufficient honey and pollen. The wax is secreted by
the hypodermal cells rather than by glands within the abdominal cavity;
the wax traverses the cuticular layer, and accumulates on its outer
surface (Carlet). According to Fritz Müller, in the stingless bees
(Trigona) which he observed, the wax-glands are situated on the back of
the abdomen, but Ihering states that in many species of Trigona and
Melipona there are also slightly developed wax-organs on the ventral
side.

It has been found that certain caterpillars secrete wax. Thus the
cells of the Tortrix of the fir (_Retinia resinella_) formed of
resin are lined with wax, as on dissolving away the resin with
alcohol, Dr. Knaggs found a slight film of wax; also a secretion of
wax has been detected in the larva of a butterfly (_Parnassius
apollo_). The bodies of certain saw-fly larvæ are covered with a
white powdery secretion, while the remarkable larva of a Selandria
is clothed with snow-white, long, flocculent, waxy masses, nearly
concealing the body (Fig. 356).

_h._ “Honey-dew” or wax-glands of Aphids

The so-called “honey-dew” of Aphids which oozes from two wart-like
tubercles or tubes situated near the end of the body, is secreted by
hypodermal unicellular glands which open into a modification of a
pore-canal, the tube itself being an outgrowth of the cuticula.

Witlaczil states that both in the “honey” tubes and in the body
beneath, the sugary matter exists in cells of the connective tissue
in the form of granules. “These large ‘sugar-cells’ in contact with
the air undergo destruction, while the sugar crystallizes into
needles, and thus each cell is transformed into a radiated
crystalline mass.”

“A muscle extends from a horseshoe-shaped place (a valve?) in the
middle of the flat terminal plate of the honey tube, through this
and down through the abdomen to the ventral surface. By this muscle
the honey tube is at times erected, and we then find, as also when
we lightly press the body of the insect, lumps of crystallized sugar
which have been expressed through the tips of the honey tubes.”
(Zool. Anzeiger, 1882, p. 241.)

[Illustration:

FIG. 356.—Wax-secreting larva of a saw-fly.
]

[Illustration:

FIG. 357.—_Lachnus strobi_, and its two “honey” warts.—Gissler
_del._
]

Busgen, after careful research, denies that this is a sugar, but
claims as the result of chemical analysis, that it is more like wax.
He observed that on reaching the air the drops issuing from the
“nectary” or “honey” tube stiffened almost instantly into a wax-like
mass, which was easily crushed between the teeth, and had no taste
at all. No sugar-like substance or urea could be detected. He
therefore concludes that the secretion in question should be
regarded as a wax-like mass, which agrees well with Witlaczil’s
anatomical observations, and confirms the statements of previous
observers. Thus, as early as 1815, Kyber stated that the Aphides
expelled an excrementitious substance through the “sap tubes.”
Burmeister states that the tubes give out a fluid which “dries
gumlike, but, so far as I have observed, has no peculiar taste.”
Réaumur, and also Kaltenbach, state that the “honey” does not issue
from the tubes, but from the anus. Lastly, Forel emphatically states
that “the two dorsal tubes of Aphides do not secrete a sweet fluid,
but a gluey wax, which is not sought by the ants. Moreover the
shield-lice and many leaf-lice have no such tubes, but yet are often
sought by ants. The drops of sugar which the ants lick up are rather
the excrement of the insects in question.” Hence the opinion first
stated by Linné, that a sweet fluid is secreted by Aphides, must be
abandoned.

On the other hand, Busgen, after careful observations, finds that
the use of the sticky, waxen secretion is in reality a protective
one, as he observed that when a larval Chrysopa rudely attacks the
Aphides, they smear its face with the sticky wax, causing at least a
momentary interruption in its attacks. He also observed that Aphides
when invaded by coccinellid larvæ set their tubes in motion and
besmear their heads and front part of the body. He thus seems to
establish the fact that these tubes secrete a protective, sticky
fluid.

_i._ Dermal glands in general

We have seen that certain of the hypodermal cells may be modified or
specialized to form secretory unicellular glands. Such are those
(trichogens) which secrete chitinous setæ, hairs, and spines, certain
setæ in some insects being hollow and containing a poison (p. 187);
others secrete wax, certain ones in Aphids “honey-dew”; in some cases
dermal glands may excrete protective, sticky, or otherwise offensive
matters, or may be depuratory, or facilitate the process of moulting.

There are other minute, unicellular, or compound dermal glands whose
function is unknown.

Dermal glands may be segmentally or serially arranged. Thus Verson has
detected a series of one or two pairs of unicellular glands near the
stigmata in each thoracic, and the first eight abdominal segments of the
silkworm (_B. mori_). In the earliest stages of growth of the
caterpillar they give out oxalate of lime, and in later stages uric
acid. They thus appear to act interchangeably with the urinary tubes, as
excretory organs. They do not, however, carry their products directly
outwards, but leave them between the hypodermis and cuticula, in order
to facilitate the sloughing off of the latter in the process of
moulting.

LITERATURE ON THE SECRETORY GLANDS

_a._ General

=Sirodot, S.= Recherches sur les sécrétions chez les Insectes. (Ann.
sc. nat., 4 Sér., Zool., 1858, x, pp. 141–189, 251–328, 12 Pls.)

=Gazagnaire, G.= Des glandes chez les Insectes. (Compt. rend. Acad.
Sc., Paris, 1886, cii, pp. 1501–1503; Annal. Soc. Ent. France, 1886,
Bull., pp. 104–106.)

=Leydig, F.= Beitrage zur Anatomie und Histiologie der Insekten. 1887.

=Hanow, Karl.= Ueber Kerfabsonderungen und ihre Benutzung im eigenen
Haushalte. (Programm des Realprogymnasiums zu Delitzsch für das
Schuljahr 1889, xc; Delitzsch, 1890, pp. 3–22.)

=Verson, E.= Di una serie di nuovi organi escretori scoperti nel
filugello. (Publ. R. Stazione Bacologica di Padova, v, 1890, pp. 30.
4 Pls.)

—— Altre cellule glandulari di origine postlarvale. (Ibid., vii, 1892,
pp. 16, 1 Pl.)

—— =ed E. Bisson=. Cellule glandulari ipostigmatiche nel _Bombyx
mori_. (Ibid., vi, 1891.)

=Borgert, H.= Die Hautdrüsen der Tracheaten. Jena, 1891, pp. 80.

=Batelli, Andrea.= Di una particolarità nell’ integumento dell’
_Aphrophora spumaria_. (Monitore Zool. Ital., 1891, Anno ii., pp.
30–32, dermal gland in last segment.)

=Koschewnikow, G. A.= On a new compound dermal gland found in the
sting of the bee. (Journal of the Zoological Section of the Society
of the Friends of Natural Science. Moscow, ii, Nos. 1, 2, 1892, p.
36. Preliminary notice. In Russian.)

=Willem, V., et H. Salbe.= Le tube ventral et les glandes céphaliques
des Sminthurus. (Ann. Soc. Ent. Belg., 1897, xli, pp. 130–132.)

=Henseval, Maurice.= Les glandes à essence der _Cossus ligniperda_.
(La Cellule, 1897, xii, pp. 19–26, 27, 29.)

—— Recherches sur l’essence der _Cossus ligniperda_. (La Cellule,
1897, xii, pp. 169–181, 183.)

_b._ Poison-glands

=Macloskie, George.= The poison-apparatus of the mosquito. (Amer.
Nat., xxii, 1888, pp. 884–888. 1 Fig. Also in Science, 1887, p.
106.)

=Beyer, Otto W.= Der Giftapparat von _Formica rufa_, ein reduziertes
Organ. (Jena. Zeitschr. f. Wissens, xxv, 1891, pp. 26–112, 2 Taf.)

=Bordas, L.= Sur l’appareil venimeux des Hyménoptères. (Comtes rend.,
cxviii, 1894, pp. 296–299 and 873–874; also Zool. Anzeiger, xvii
Jahrg., 1894, pp. 385–387, Figs.)

—— Appareil glandulaire des Hyménoptères. (Glandes salivaires; Tubes
de Malpighi et glandes venimeuses.) Paris, 1894, pp. 362, 11 Pls.

—— Description anatomique et étude histologique des glandes à venin
des insectes Hyménoptères. Paris, 1897, pp. 53, 2 Pls.

See Forel (p. 186); also the standard authors, Kolbe, etc.; also Locy
(Amer. Nat., xviii, 1884, p. 355), Nagel (p. 324), Fenger.

_c._ Wax-glands

=Brandt und Ratzeburg.= Medicinische Zoologie, ii, 1830, p. 179. Taf.
xxv, Fig. 18.

=Treviranus, George R.= Ueber die Bereitung des Wachses durch die
Bienen. (Zeitschrift für Physiologie, etc., iii, 1832, pp. 62, 225.)

=Dufour, L.= Note anatomique sur la question de la production de la
cire des abeilles. (Comptes rend. Acad. Sc., Paris, 1843, xvii, pp.
809–813, 1248–1253; Revue Zool. 1843, l’Institut, 1843, xi.)

=Dujardin, F.= Mémoire sur l’étude microscopique de la cire, etc.
(Ann. Sc. Nat., xii, 1849, pp. 250–259.)

=Tarzione-Tozzetti, H.= Studii sulle cocciniglie. Milano, 1867, 7 Pls.

—— Sur la cire qu’on peut obtenir de la cochenille du figuier (_Coccus
caricæ_). (Comptes rend. Acad. Sc., Paris, lxv, 1867, pp. 246–247.)

=Claus, C.= Ueber die wachsbereitenden Hautdrüsen der Insekten.
(Sitzungsber. Gesells. z. Beförd. d. Gesammt. Naturw. zu Marburg,
June, 1867, No. 8, pp. 65–72.)

=Witlaczil, E.= Die Anatomie der Psylliden. (Zeitschr. wissens. Zool.,
xlii, 1885, pp. 582–586.)

—— Zur Morphologie und Anatomie der Cocciden. (Zeitschr. wissens.
Zool., xliii, 1886, pp. 149–174, 1 Taf.)

=Ihering, H. von.= Der Stachel der Meliponen. (Ent. Nachrichten, xii
Jahrg., 1886, p. 185.)

=Carlet, G.= Sur les organes sécréteurs et la sécrétion de la cire
chez l’Abeille. (Comptes rendus, cx, pp. 361–363, 1890.)

—— La cire et ses organes sécréteurs. (Le Naturaliste, 1890, pp.
149–151, 2 Figs.)

Also the works of Siebold, Cheshire, Kolbe, Howard and Marlatt (Bull.,
3, N. S., Div. Ent. U. S. Dept. Agr., 1896, p. 40), Knaggs, Berlese.

_d._ Wax-like glands of Aphides

=Huber et Forel, A.= Études myrmécologiques, 1875.

=Witlaczil, E.= Zur Anatomie der Aphiden. (Zool. Anzeiger, v Jahrg.,
1882, pp. 239–241; Arbeiten a. d. Zool. Institut der Univ. Wien.,
iv, 1882, pp. 397–441, 3 Taf.)

=Busgen, M. J.= Der Honigtau. Biol. Studien an Pflanzen u.
Pflanzenlause. (Jena. Zeitschrift, xxv, 1891, pp. 339–428.)

DEFENSIVE OR REPUGNATORIAL SCENT-GLANDS

While these eversible glands are not found in marine or aquatic
arthropods such as Crustacea or Merostomata (Limulus), they are often
present in the air-breathing forms, especially insects. In the winged
insects they are of frequent occurrence, existing under great variety of
form, varying greatly in position, and appearing usually to be in
immediate relation with their active volant habits. Their presence is in
direct adaptation to the needs and habits of their possessors, and being
repellent, warning, or defensive structures, the odors they secrete
being often exceedingly nauseous, they appear to have been called into
existence in direct response to their biological environment. The fact
that these singular organs do not exist in marine or aquatic Crustacea
suggests that the air-breathing, aërial, or volant insects by these
eversible glands, usually in the form of simple evaginable hypodermic
pouches, are enabled to protect themselves by emitting an infinitesimal
amount of an offensively odorous fluid or ether-like spray which charges
the air throughout an extent of territory which may be practically
illimitable to the senses of their enemies. The principle is the same as
in the mephitic sulphuretted oil ejected by the skunks, the slight
quantity these creatures give out readily mixing with and charging the
atmosphere within a radius of many miles of what we may call the centre
of distribution.

As is now well known, the very delicate, attenuated highly volatile
odors exhaled are perceived by insects with extreme ease and rapidity,
the degree of sensitiveness to such scents being enormously greater than
in vertebrates, their organs of sense being developed in a corresponding
degree. Professors Fischer and Penzoldt, of Erlangen, have recently
established the fact that the sense of smell is by far the most delicate
of the senses. They find that the olfactory nerve is able to detect the
presence of 1⁄2,760,000,000 of a grain of mercaptan.[58] The smallest
particle of matter that can be detected by the eye is sodium, when
observed by the spectroscope, and this particle is 250 times coarser
than the particle of mercaptan which can be detected by the human nose.

In those Arachnida which are provided with poison-glands, these
scent-glands are absent, but in certain Acarina and Linguatulidæ,
which have no poison-glands, there are various oil-glands, stigmatic
glands, as well as scent-glands, and in seizing a Thelyphonus with
the forceps we have observed it to send out from each side of the
body a jet of offensive spray.

We not infrequently find in myriopods (Polydesmidæ, Julidæ, and
Glomeris) repugnatorial or the so-called cyanogenic glands, which
are either paired, opening on the sides of the body, or form a
single row along the median line of the under side of the body.
Leidy describes and figures the spherical glands of _Julus
marginatus_, of which there are 50 pairs. These glands have been
regarded as modified nephridia, but are more probably coxal glands,
and the homologues of the parapodial glands of annelid worms.

[Illustration:

FIG. 358.—Sternite of _Machilis maritima_, with the pair of coxal sacs
(_cb_) on the right side everted; _hs_, coxal appendages; _m_,
retractor muscles.—After Oudemans, from Lang.
]

=Eversible coxal glands.=—True coxal glands occur in _Scolopendrella
immaculata_ on the 2d to 11th segment, on the inner side of the base of
the legs (Fig. 15, _c.g._). Homologous glands also occur in the same
position in _Campodea staphylinus_ (also in _C. cookei_ and _C.
mexicana_) on the 1st to 8th abdominal segments, and Oudemans has
described a pair of eversible sacs on each side of segments one to seven
of Machilis. These eversible sacs in the synapterous insects are
evidently modified coxal glands, and are probably repugnatorial as well
as respiratory in function.

The apparatus consists of an eversible gland, composed of hypodermic
cells, usually retracted by a slender muscle and with an efferent
passage, but the glands vary greatly in shape and structure in different
insects. In some cases these fœtid glands appear not to be the
homologues of the coxal glands, but simply dermal glands.

These repugnatorial glands are of not infrequent occurrence in the lower
or more generalized winged insects, and in situation and appearance are
evidently the homologues of the coxal glands of the Symphyla and
Synaptera.

_Fœtid glands of Orthoptera._—In the ear-wigs (Forficula and Chelidura)
Meinert has detected a pair of what he calls fœtid glands at the
posterior margin of the dorsal plates of the 2d and 3d abdominal
segments.

Vosseler also describes the same glands as consisting of a retort-shaped
sac, in whose walls are numerous small hypodermal cells and large single
glandular cells provided with an efferent passage, the fluid being
forced out by the pressure of the dermal muscles, one acting specially
to retract the gland. The creature can squirt to a distance of 5 and
even 10 cm. (4 inches) a yellowish-brown liquid or emulsion with the
odor of a mixture of carbolic acid and creosote.

The large eversible dorsal glands of the Blattidæ, since they contain
numerous hairs, which, when everted, are fan-like or like tufts, serve,
as in the spraying or scent apparatus, to disseminate the odor, and
might be classified with the alluring unicellular scent-glands or
_duftapparat_ of other insects, as they are by some authors; but as the
glands are large and compound they may prove to be the homologues of the
coxal glands rather than of the dermal glands.

Evaginable organs in the Blattids were first observed by Gerstæcker in
both sexes of Corydia; they are yellowish white, covered with hairs, and
are thrust out from between the dorsal and ventral plates of the 1st and
2d abdominal segments.

[Illustration:

_Fig. 359._—Under side of end of Aphlebia, showing the two eversible
sacs; _V-X_, five last abdominal segments; _A_, portion showing the
hairs; _B_, showing origin of a hair in its follicle.—After Krauss.
]

In the cockroach (_P. orientalis_) Minchin detected two pouch-like
invaginations of the cuticle, lying close on each side of the middle
line of the body between the 5th and 6th tergites of the abdomen. They
are lined by a continuation of the cuticle, which forms, within the
pouches, numerous stiff, branched, finely pointed bristles, beneath
which are a number of glandular epithelial cells. In the male nymph of
_P. decorata_ he also found beside these glandular pouches “an
additional gland, opening by a tubular duct under the intersegmental
membrane between the 5th and 6th terga above the glandular pouch of each
side, and extending forward into the body cavity. The gland and its duct
are proliferations of the hypodermis, and there is no invagination of
the cuticle.” These eversible glands are most complicated in
_Phyllodromia germanica_. While it is absent in the female, in the male
it is relatively of enormous size, extending over the 6th and 7th
somites, as well as projecting far into the body cavity (Minchin). Haase
states that these glands become everted by blood-pressure and give out
the well-known disagreeable smell of these insects. He states that in
the male of _P. germanica_ the dorsal glands in the 6th and 7th
abdominal segments are without hairs and produce an oily secretion; they
function as odoriferous organs in sexual union.

In the male of another Blattid (_Aphlebia bivittata_) of the Canary
Islands, Krauss has detected two yellowish dorsal sacs 1.5 mm. in
length, opening out on the 7th abdominal segment, and filled full of
long yellowish hairs, the ends directed towards the opening, where they
form a thick tuft. These eversible glands lined with hairs appear to be
closely similar to the long slender eversible hairy appendages or scent
organs of certain Arctian and Syntomid moths. (Fig. 359.)

[Illustration:

FIG. 360.—External flaps (_gl_) of glands of Platyzosteria.
]

We have found the external median wart with lateral lids or flaps in
between the 5th and 6th tergites of _Platyzosteria ingens_ Scudder, a
large wingless Blattid living under the leaf scars of the cocoanut tree
in Southern Florida (Fig. 360), but were unable to detect them in
Polyzosteria or in the common Blabera of Cuba, or in another genus from
Cordova, Mexico.

In another group of Orthoptera, the Phasmidæ, occur a pair of dorsal
prothoracic glands, each opening by a pore and present in both sexes. In
the walking-stick, _Anisomorpha buprestoides_, ♂ and ♀, these openings
are situated on each side of the prothorax at its upper anterior
extremity, situated at the bottom of a large deep pit. When seized it
discharged a “milky white fluid from the pores of the thorax, diffusing
a strong odor, in a great measure like that of the common Gnaphalium or
life everlasting” (Peale in Say’s American Entomology, i, p. 84). Boll
states that the females when captured “spurt from the prothorax,
somewhat after the manner of bombardier beetles, a strong vapor, which
slightly burnt the skin; when the females were seized by the males a
thick fluid oozed from the same spot.” Scudder describes these glands in
another Phasmid (_Autolyca pallidicornis_) as two straight, flattened,
ribbon-like bodies, with thick walls, broadly rounded at the end, lying
side by side and extending to the hinder end of the mesothorax. In
_Anisomorpha buprestoides_ the glands are of the same size and shape
(Scudder). In _Diapheromera femorata_ the repugnatorial foramina are
very minute, and the apparatus within consists of a pair of small
obovate or subfusiform sacs, one on each side of the prothorax, about 1
mm. in length, with a short and very slender duct opening externally at
the bottom of the pit (Scudder).

In the Mantidæ these seem to be genuine coxal glands, as there is a pair
situated between the coxæ of the first pair of legs. An evaginable organ
like a wart, with a glandular appearance, occurs on the hind femora of
the Acrydiidæ in a furrow on the under side, into which the tibia fits,
about one-fourth from the base (Psyche, iii, p. 32).

In the male cricket, the anal odoriferous glands are small lobes opening
into a reservoir on each side of the rectum (Dufour). Homologous glands
also occur in the Coleoptera (Fig. 302, _l_ and 317, _s_).

[Illustration:

FIG. 361.—Glands (_g_) of Lachnus; _h_, “honey” wart.—Gissler _del._
]

Most Hemiptera or bugs send out a fœtid or nauseous odor due to a fluid
secreted by a single or double yellow or red pear-shaped gland, situated
in the middle of the mesothoracic segment, and opening between the
hinder or third pair of coxæ. In Belostoma Leidy describes these glands
as consisting of two rather long cœcal tubes situated in the metathorax,
beneath the other viscera, extending backwards into the abdomen, and
opening between the coxæ of the third pair of legs. Locy states that the
smell arising from these glands is pleasant, resembling that of well
ripened pears or bananas. Other bugs, moreover, emit an agreeable odor,
that of Syromastes resembling that of a fine bergamot pear. (Siebold.)
The fluid given out by the European fire-bug (_Pyrrhocoris apterus_) has
a sweetish smell, like ether. In the nymph there are three pairs of
dorsal glands, on abdominal segments 2–5, which are atrophied in the
mature insect. In the bed-bug, the nymph has three odoriferous glands
each with paired openings in the three basal abdominal segments
respectively, and situated on the median dorsal line, being arranged
transversely at the edge of the tergites; but after the last moult these
are aborted, and replaced by the sternal metathoracic glands (Künckel).
Gissler has detected a pair of glands in _Lachnus strobi_ (Fig. 361).

=Anal glands of beetles.=—Certain beetles are endowed with eversible
repugnatorial glands. _Eleodes gigantea_ and _E. dentipes_ of both sexes
are said by Gissler to possess these glands. When teased “they stand on
their anterior and middle legs, holding the abdomen high up and spurting
the contents of the glands right and left.” The glands (Fig. 366, 1) are
two reddish brown, somewhat bilobed sacs, and extend from the base of
the last up to the middle of the 2d abdominal segment, with an average
length of 6.5 mm. The liquid stains the human skin, has an acid
reaction, with a peculiar, “intensely penetrant odor, causing the eye to
lachrymate. It is soluble in water, alcohol, and ether. Boiled with
concentrated sulphuric acid and alcohol an ethereal aromatic vapor is
produced, indicating the presence of one or more organic acids, though
neither formic or acetic acid could be detected.” Williston has observed
the same habits in seven other species of Eleodes, all ejecting a
pungent vile-smelling liquid, one species (_E. longicollis_) ejecting a
stream of fluid from the anal gland, backwards sometimes to the distance
of 10 cm. or more, and he regards these beetles as “the veritable skunks
of their order.” Leidy briefly describes the odoriferous glands of _Upis
pennsylvanica_.

The anal glands consist, according to Meckel and also Dufour, of two
long, simple, flexuous cœca with reservoirs having two short excretory
ducts situated near the anus (Siebold).

Glands like those of Eleodes found in _Blaps mortisaga_ are described in
detail by Gilson (Fig. 366, 2). They form two pouches or cuticular
invaginations situated in the end of the abdomen on the sides of the end
of the intestine and unite on the median line underneath the genital
organs, forming a very short tube with a chitinous wall, continuous with
the cuticle of the last abdominal segment. Into each pouch open a large
number of fine slender lobules varying in shape, giving a villous aspect
to the surface. These lobules are composed of as many as fifty
unicellular glands, each of which is composed of four parts: (1) A
radiated vesicle, (2) a central sac, giving rise (3) to a fine excretory
tube, and (4) a sheath near the origin of the excretory tube. These are
all modifications of the cytoplasm of the cell with its reticulum; the
nucleus with its chromosomes is also present, but situated on one side
of the central sac. The fine excretory tubules form a bundle passing
down into the mouth of each lobule.

Similar glands, though usually smaller, which have not been
carefully examined, occur in Carabus (Fig. 300, 3) and Cychrus,
which eject from the vent a disagreeable fluid containing butyric
acid (Pelouse). The bombardier beetle Brachinus, with its anal
glands, ejects a jet of bluish vapor accompanied with a considerable
explosion, which colors the human skin rust red; it is caustic,
smells like nitrous acid, and turns blue paper red. Westwood states
that individuals of a large South American Brachinus on being seized
“immediately began to play off their artillery, burning and staining
the flesh to such a degree that only a few specimens could be
captured with the naked hand, leaving a mark which remained for a
considerable time.” The fluid ejected by another species, in
Tripoli, blackened the fingers of the collector. “It is neither
alkaline nor acid, and it is soluble in water and in alcohol.”
(Kirby and Spence, iv, p. 149.)

Species of other genera (Agonum, Pheropsophus, Galerita, Helluo,
Paussus, Ozæna) are also bombardiers, though less decidedly so than
Brachinus. A Paussid beetle (Cerapterus) ejects explosively a fluid
containing free iodine (Loman), while Staphylinus, Stenus, Ocypus
olens, Lacon, etc., have similar anal fœtid glands, the liquid being
more or less corrosive. The secretion of _Mormolyce phyllodes_ is so
corrosive that it is said to paralyze the fingers for 24 hours
after. (Cuénot.)

[Illustration:

FIG. 362.—Median section through the femoro-tibial joint of leg of
Coccinella, showing at _o_ the opening through which the blood oozes
out; _f_, femur; _t_, tibia; _e_, extensor muscle of the tibia: _s_,
sinew of the same; at _ch_, chitinized; _h_, articular membrane;
_v_, tibial process.—After Lutz.
]

The two pairs of remarkably large, soft, eversible, forked,
orange-yellow glands of the European genus Malachius, are thrust out
from the side of the 1st and the 3d thoracic segments. They are everted
by blood-pressure, and retracted by muscles. The larva of _Hydrophilus
piceus_ ejects by the anus a black, fœtid fluid.

Claus has shown that the larva of _Lina populi_ and other Chrysomelidæ
possess numerous minute, eversible glands in each of the warts on the
upper surface of the body, each gland containing a whitish, repellent
fluid smelling like the oil of bitter almonds and containing salicylic
acid derived from its food-plant, which issues as pearl-like drops.
Candèze thinks the fluid may contain prussic acid. The fluid is secreted
by a variable number of glandular cells, each provided with an efferent
duct. The larvæ of saw-flies, notably _Cimbex americana_, also eject
droplets of a clear fluid from non-evaginable glands situated near each
stigma (Chlolodkovsky).

=The blood as a repellent fluid.=—In this connection it may be mentioned
that though there are no special glands present, many beetles emit drops
of blood from the femoro-tibial joints of their legs as a means of
defence. Such are the oil-beetles (Meloë), Cantharis, Lytta. The
cantharadine secreted by these beetles, according to Beauregard, is an
efficient means of defence, as birds, reptiles, and carnivorous insects
will not usually attack them. This substance is formed in the blood and
also in the genital organs, and is so extremely caustic that scavenger
insects which feed upon their dead bodies will leave untouched the parts
containing cantharadine, and if May-beetles or mole-crickets are washed
with the blood of Meloë or with cantharidate of potassa, it will for
several days render them safe from the attacks of the carabids which
usually prey upon them. The eggs even after deposition are strongly
vesicant, and are thus free from the attacks of egg-eating insects
(Cuénot). The Coccinellidæ are also protected by a yellow, mucilaginous,
disagreeable fluid oozing out of the ends of the femora; in our common,
two-spotted lady-bird (_C. bipunctata_) the yellow fluid is
disagreeable, smelling like opium. Lutz has found that the blood in
Coccinellidæ passes out through a minute opening situated at the end of
each femur (Fig. 362). The blood is very repellent to insectivorous
animals.

The Dyticidæ eject from the anus a colorless, disagreeable fluid, while
these beetles, and especially the Gyrinidæ, when captured send out a
milky fluid which appears to issue from the joints of the body. The
Silphidæ throw out both from the mouth and vent a fœtid liquid with an
ammoniacal odor. They possess but a single anal gland, the reservoir
opening on one side of the rectum (Dufour).

Other malodorous insects have not yet been investigated; such are the
very persistent odors of lace-winged flies (Chrysopa).

More agreeable secretions, but probably formed by similar glands, is the
odor of rose or hyacinth given out by Cicindelæ, or the rose fragrance
exhaled by the European _Aromia moschata_.

=Eversible glands of caddis-worms and caterpillars.=—Gilson, while
investigating the segmentally disposed thoracic glands of larval
Trichoptera, has found in the larva of _Limnophilus flavicornis_ that
the sternal prothoracic tubercle gives exit to an underlying tubular
gland. In _Phryganea grandis_ each thoracic sternum affords an exit to
an eversible gland. Many caterpillars, as our subjoined list will show,
are very well protected by eversible repugnatorial glands situated
either in the under or upper side of the body. Since the time of De Geer
(1750) the fork-tailed larva of Cerura has been known to throw out a
secretion, which was described by Bonnet in 1755 as a true acid, sharp,
sour, and biting. This spraying apparatus in _Cerura (Harpyia) vinula_
has been well described by Klemensiewicz (Fig. 366, 4), though Rengger
in 1817 noticed the general form of the secretory sac, and that it opens
out in two muscular eversible tubes, out of which the secretion is
ejected.

The fork-tailed larva of _Macrurocampa marthesia_, which is much like
that of Cerura, when teased sends out a jet of spray to the distance of
nearly an inch from each side of the neck. While examining the very
gayly-colored and heavily-spined caterpillars of _Schizura concinna_ we
observed that when a fully-grown one was roughly seized with the forceps
or fingers it sent out a shower of spray from each side of the
prothoracic segment, exactly like that of Cerura and Macrurocampa.

In the European _Cerura vinula_ the apparatus consists of a single sac,
which opens by a narrow transverse slit on the under side of the neck,
out of which is rapidly everted four lateral tubes, two on each side
(Fig. 366, 4, _t_), which are withdrawn within the opening by the
contraction of several fine muscles. The apparatus in the American _C.
multiscripta_ is as in the European _C. vinula_. In a living specimen
the large secretory sac was seen to be of the same size and shape as in
Macrurocampa, and of the color of raw silk. The sac when distended
extends back to a little behind the middle pair of legs, and is nearly
two-thirds as wide as the body. The caterpillar sent out the fluid when
handled, but we could not make it spray.

In the larva of _Macrurocampa marthesia_ the cervical or secretory gland
(Fig. 366, 5) is situated in the 1st and 2d thoracic segments, extending
to the hinder edge of the latter and lying between the nervous cord and
the œsophagus and proventriculus, and when empty the bulk of it lies a
little to one side of the median line of the body. It is partly held in
place by small tracheæ, one quite large branch being sent to it from
near the prothoracic spiracle. The short, large duct, leading from it to
the transverse opening in the membrane between the head and prothoracic
segment, is a little narrower than this opening, and is kept distended
by tænidia, or a series of short, spiral threads which are pale, not
honey-yellowish, in color. This duct lies on one side of the prothoracic
ganglion, resting just under the commissures passing up to the brain; it
is also situated between the two silk ducts.

The very distensible sac (Fig. 366, 5) is rendered elastic by a curious
arrangement of the cuticle, the tænidia of the duct itself being
represented by very thickly-scattered, irregular, separate, sinuous,
chitinous ridges, which stand up from the cuticular lining of the wall
of the sac (Fig. 366, 6). The secretory cells of the walls of this sac
in _Cerura vinula_ are said by Klemensiewicz to be large hexagonal
cells, resembling those of silk-glands, having like them large branched
nuclei.

The fluid thrown out is said by Poulton to be formic acid; it causes
violent effervescence when allowed to fall upon sodium-bicarbonate, and
colors blue litmus paper red. It also appears from the researches of
Latter that these creatures in the imago state secrete free potassium
hydroxid, a substance for the first time known to exist in the animal
kingdom.

In the caterpillar of _Astyanax archippus_ (_Limenitis disippus_) a
dark, bladder-like sac is everted, but the lateral tubes appear to be
wanting, and no spray is sent out; it occurs in the larvæ of many
Nymphalidæ and other butterflies and moths.

These glands are functionally active in Perophora, but obsolete (at
least the external openings) in Lacosoma.

=The osmeterium in Papilio larvæ.=—The caterpillars of the swallowtailed
butterflies (Papilio, Doritis, and Thais), as is well known, when
irritated thrust out from a transverse slit on the upper part of the
prothoracic segment a large orange-yellow V-shaped fleshy tubular
process (the osmeterium), from which is diffused a more or less
melonlike but disagreeable, in some cases insufferable, odor; the
secretion is acid and reddens litmus paper. The mechanism has been
described and figured by Klemensiewicz.

When at rest, or retracted, the osmeterium lies in the upper part of the
body in the three thoracic segments, and is crossed obliquely by several
muscular bundles attached to the walls of the body, and by the action of
these muscles the evagination of the osmeterium is strongly promoted.
After eversion the tubes are slowly retracted by two slender muscles
inserted at the end of each fork or tube, and arising from the sides of
the 3d segment behind the head, crossing each other in the median line
(Fig. 366, 7 _r.m._). The secretion is formed by an oval mass of
glandular cells at the base of the forks; in the glandular mass is a
furrow-like depression about which the secretory cells are grouped. The
secretion collects in very fine drops on the side of each furrow
opposite the glandular cells.

According to C. D. Ash the larva of an Australian Notodontid (_Danima
banksii_ Lewin) protrudes from the under side of the prothoracic segment
a Y-shaped red organ like that of Papilio; no fluid or odor is given
out.

=Dorsal and lateral eversible metameric sacs in other larvæ.=—The showy
caterpillars of Orgyia and its allies have a conspicuous coral-red
tubercle on the back of the 6th and also the 7th abdominal segment,
which on irritation are elongated, the end of the tubercles being
eversible. When at rest the summit is crateriform, but on eversion the
end becomes rounded and conical. These osmeteria are everted by blood
pressure, and retracted by a muscle. Fig. 366, 9, represents a section
of an osmeterium of _Orgyia leucostigma_ when retracted by the muscle
(_m_); at the bottom of the crater are the secreting or glandular cells
(_gc_), being modified hypodermal cells. These doubtless serve as
terrifying organs to ichneumons and other insect enemies, and though we
have been unable to detect any odor emanating from the tubercles, yet
possibly they give out a scent perceived by and disagreeable to their
insect assailants.

[Illustration:

FIG. 363.—Freshly hatched larva of _Hyperchiria io_, with its two
pairs of eversible glands (_g_).
]

[Illustration:

FIG. 364.—Young larva of _Megalopyge crispata_, enlarged, showing the
seven pairs of lateral processes (_lp_): _sp_, spiracle; _abl′_,
_abl^6_, six pairs of abdominal legs besides the anal pair.
]

In the Hemileucidæ there is a pair of lateral osmeteria, on the 1st and
on the 7th abdominal segments, which however, are not highly colored
(Figs. 363, 366, 10). In Megalopyge (Lagoa, Fig. 364) there is a lateral
row of singular pale permanently everted processes which appear to be
the homologues of the osmeteria of larvæ of other lepidopterous
families. As these are repeated on seven segments, their metameric
arrangement is obvious. The relation of these curious glands to the
viscera is seen in Fig. 297, _lgp_, and their minute structure in Fig.
365.

At _A_, the lumen (_l_) is a deep narrow cavity, with the secretion
(_secr._), collected at the mouth of the cavity, composed of a thin,
mucus-like, coagulated fluid, containing granules of varying degrees
of fineness, which take the stain readily. Outside of these are
collected fine nuclei (_bc_), stained dark, and enveloped in a
slight, transparent, pale, protoplasmic envelope, which may be blood
corpuscles. The glandular cells themselves are simply modified
hypodermal cells, as seen at _C_. In some of the nuclei, indistinct
nucleoli are seen, and deeply stained granules, especially around
the periphery of the nuclei. At _B_ is represented a section on one
side of the middle, but still showing the spacious lumen. In the
section represented by _C_, the knife passed through the process
still nearer the outer edge, and near the base; at _C^1_, three of
the glandular cells, with their large, deeply stained nuclei, are
drawn. A transverse section at _D_ shows the large lumen or cavity
(_l_).

As to the function and homologies of these structures, it is
difficult to decide. We have never noticed that they give off any
odor, though they may prove to be repugnatorial; they are not
visible in the fully grown, living insect, being concealed by the
long, dense hairs clothing the body; they are not spraying organs,
as they are imperforate at the end, not ending as the lateral,
eversible glands of _Hyperchiria io_, etc., in a crateriform
orifice.

They may be permanently everted glands, or osmeteria, which have, by
disuse, lost their power of retraction and their crateriform
opening, as well as the power of secreting a malodorous fluid.

[Illustration:

FIG. 365.—Section of lateral processes of larva of Megalopyge.
]

In certain of the butterflies, the Heliconidæ (Colænis, Heliconius,
Euides, and Dione), there is thrust from the end of the abdomen a pair
of large, irregular, rounded, eversible glands, which give out a
disagreeable odor, and are consequently repellent, and which seem to be
the homologues of the odoriferous glands of other butterflies.

The large, soft, rounded, eversible glands, looking like puff-balls
or a rounded pudding (Fig. 366, 12), are everted, when the
butterflies are roughly seized, from the dorsal side of the
penultimate segment of the abdomen. The males possess two smaller
tubercles on the inside of the anal claspers or lobes. Müller also
detected, in the females of various species of the Heliconidæ
enumerated above, a pair of club-shaped processes like the balancers
of flies, which are thrust out on each side of and under the
odoriferous puff-balls of the hinder edge of the penultimate segment
(Fig. 366, 13). The club or head is armed with hairs or bristles,
which, in Heliconius, are like the scales of a butterfly.

[Illustration:

FIG. 366.—Scent-glands of insects: 1. Anal eversible glands of
Eleodes.—After Gissler. 2. Anal eversible glands of Blaps.—After
Gilson. 3. Anal glands (_agl_) of _Carabus hortensis_: _rs_,
reservoir; _d_, excretory duct; _i_, intestine; _r_, rectum.—After
Kolbe. 4. Prothoracic spraying apparatus of _Cerura vinula_: _gl_,
the gland; _d_, its duct, with tænidia; _t_, the spraying tubes;
_m_, muscles; _rm_, retractor muscles.—After Klemensiewicz. 5. The
thoracic glandular sac of _Macrurocampa marthesia_: _gl_, the
glandular sac; _d_, its duct; _e_, peritracheal epithelium; _t_, the
spiral threads or tænidia. 6. Irregular separate masses of chitinous
ridges on the cuticular lining of the wall of the sacs of
_Macrurocampa marthesia_. 7. Osmeterium (_os_) of the larva of
_Papilio machaon_ at rest: _rm_, the retractor muscles at the ends;
_m_, the numerous oblique muscles; _dm_, dorsal longitudinal
muscles; _t_, trachea; _oe_, œsophagus; _gang_, brain; 1, head; 2,
3, 4. thoracic segments. 8. Osmeterium (_os_) of one side, enlarged:
_g_, glandular portion at the base; _d_, depressions in the cuticula
of the glandular portion; _t_, trachea.—This and Fig. 7 after
Klemensiewicz. 9. Eversible dorsal glands (_ev. gl_) of larva of
_Orgyia leucostigma_ in Stage II: _gc_, glandular cells at bottom of
the crater-like depression; _m_, retractor muscle; _p_, poison
gland-cells of the root of the seta (_s_); _c_, cuticula; _hyp_,
hypodermis; _A_, portion of the cuticle and hypodermis enlarged. 10.
Lateral eversible gland of _Hyperchiria io_, Stage II: _rm_,
retractor muscle; _oen_, œnocytes. 11. The same as Fig. 10, but
representing a section through one side of the eversible gland. 12.
_A_, end of body of _Colænis julia_; _ev_, eversible anal gland;
_oa_, odoriferous appendages; _B_, the same in _Heliconius
apseudes_, side view; _C_, odoriferous appendages of _Colænis dido_
in fresh condition; _D_, tested with alcohol and benzine. 13.
Odoriferous appendages of _Heliconius eucrate_, head cleansed.—Figs.
12, 13, after F. Müller. 14. Odoriferous glands (_ogl_) in the pupa
of _Vanessa io_: _r_, rectum; _h_, the folds of hypodermis which
forms the terminal papilla of the abdomen; _ov_, oviduct.—After
Jackson.
]

In the caterpillars of certain blue butterflies (Lycænidæ) is an
internal osmeterium, being a very minute sac which is everted from a
transverse slit on the top of the 7th abdominal segment. Its function is
quite the opposite of those of the caterpillars of other families, since
the sac exudes a sweet fluid very attractive to ants, which may be
diffused more widely by the delicate spinulose bristles crowning the
summit. W. H. Edwards states that in several species of Lycæna, besides
that on the 7th abdominal segment, there is on the 8th segment a pair of
minute dorsal evaginable tubercles.

A pair of small ramose odoriferous glands are said by Siebold, who
regarded them as alluring glands, to occur in Argynnis, Melitæa, and
Zygæna, to be situated near the orifice of the oviduct, and Scudder has
detected them near the anus of the female pupa of _Danais archippus_.
The appearance of the odoriferous glands in the pupa of _Vanessa io_ is
well shown by Jackson (Fig. 366, 14). They develop as two tubular
ingrowths of the hypodermis, perfectly distinct one from the other, each
having its own separate aperture to the exterior. In Fig. 366, 14 the
condition of parts is nearly as in the imago, the glands being situated
below the rectum and opening of the oviduct. In both sexes of another
Brazilian butterfly (_Didonis biblis_) on the median line of the abdomen
between the 4th and 5th segments are two roundish vesicles covered with
short gray hairs, which emit a disagreeable smell.

It is possible that the dark-green fluid in Parnassius, secreted by
an evaginable gland, and which is moulded into shape by the
scimetar-shaped peraplast (Scudder), is formed by the homologues of
the anal glands of other butterflies.

Distribution of repugnatorial or alluring scent-glands in insects[59]

_A._ LARVAL INSECTS

_a._ _Each thoracic segment; sternal. Phryganea grandis._

_b._ _Prothoracic, sternal, discharging a lateral jet of spray; with a
single large internal sack._

LEPIDOPTERA

Family TINEIDÆ

_Hyponomeuta evonymella._

Family NOCTUIDÆ

_Bryophila_, _Cucullia formosa_, _C. scrophulariæ_, _Habrostola_,
_Cleophana linariæ,_ Catocala (sp.), _Aporia cratægæ_, _Aplecta
nebulosa_, _Leucania staminea_, _L. hispanica_, _L. nonagrioides_,
_Plusia gamma_.

Family NOTODONTIDÆ

_Pheosia rimosa_, _Schizura concinna_, _Danima Banksii_ (Australia),
_Macrurocampa marthesia_, _Heterocampa pulverea_, _Cerura vinula_, _C.
furcula_, _C. borealis_, _C. multiscripta._

Family NYMPHALIDÆ

Probably all the species.

_c._ _Prothoracic, dorsal; sending out a_ V-_shaped odoriferous organ_
(osmeterium).

Family PAPILIONIDÆ

All the species as a rule.

_d._ _Thoracic sternal, evaginable glands._

Family PEROPHORIDÆ

_Lacosoma chirodota_, _Perophora melsheimerii_.

Family NOLIDÆ

In three, and probably in all the species of Nola.

_e._ _Lateral, abdominal, non-eversible glands, one near each spiracle,
emitting a clear fluid._

Family TENTHREDINIDÆ

_Cræsus septentrionalis_, _C. varus_, _Cimbex americana_, _C. betulæ,
Trichiosoma_.

_f._ _Lateral, abdominal, partly eversible glands emitting neither
moisture nor odor, but flesh-colored._

Family _Tineidæ_

Phyllocnistis? (eight pairs.)

Family HEMILEUCIDÆ

_Hyperchiria io_ (two pairs, viz. on 1st and 7th segments), _H._ sp.
(Mexico), _Hemileuca yavapai_, _pamina_, _H. maia_, _H. artemis_,
_Pseudohazis eglanterina_.

_g._ _Lateral, abdominal, permanently everted, metameric glands, not
known to secrete a fluid, nor to be odoriferous._

Family MEGALOPYGIDÆ

_Megalopyge crispata._

_h._ _Medio-dorsal, partly eversible glands, emitting a spray of liquid
but no odor(?), and colored coral-red or orange-yellow (P. auriflua),
but usually in the European species yellowish._

Family LIPARIDÆ

All the species except those of Demas.

_i._ _A single, median, abdominal, dorsal gland, emitting a fluid
attractive to ants, on 7th segment; with a pair of minute, index glands
on the 8th segment._

Family LYCÆNIDÆ

All the species.

_j._ _Protrusile organs near the anus._

_Myrmeleon_ larva (Hagen? Dimmock).

_B._ NYMPH OF HETEROMETABOLOUS INSECTS

_a._ _Paired, dorsal glands, on abdominal segments 1, 2, and 3._

_Cimex lectularius_ (Künckel).

_b._ _The same on abdominal segment 5._

_Lachnus strobi._

_C._ PUPA OF CERTAIN BOMBYCES

_At anterior end of certain pupæ, internal glands to moisten threads of
the cocoon for exit of moth._

_D._ ADULT INSECTS

_a._ _Occurring on the prothorax only; strongly repugnatorial, best
developed in_ ♂.

_Anisomorpha buprestoides_, _Autolyca pallidicornis_, _Phasma putidum_,
_Phyllium_ (sp.), _Heteropteryx_ (sp.), _Diapheromera femoratum_
(probably in all the species of the family), _Mantis carolina_.

_b._ _Occurring on the pro- and mesothorax, and on the middle of the
abdomen, orange-yellow, fleshy tubercles or evaginations._

_Malachius bipustulatus_, _Anthocomus equestris_, _Evæus thoracicus_.

_c._ _Segmental, eversible glands, homologues of the coxal glands of
other Arthropods, occurring on all, or nearly all, the abdominal
segments._

_Scolopendrella immaculata_ (coxal glands on 3d to 11th pair of legs),
_Campodea staphylinus_ (a pair of coxal glands on 1st to 8th abdominal
segments), _Machilis maritima_ (eversible, coxal glands on segments
1–7).

_d._ _Occurring in the abdomen._

_d^1._ _In the two first abdominal segments._

_Corydia carunculigera_ ♂ and ♀.

_d^2._ _Alluring (?) organs situated on the dorsal side of the abdomen,
in the 6th, or 6th and 7th, abdominal segment._

_Periplaneta americana_ ♂, _P. orientalis_ (nymph), _P. decorata_ ♂
(nymph), _Ectoblatta germanica_ ♂, _Ectobia lapponica_ ♂, _Phyllodromia_
♂, _Aphlebia bivittata_ ♂, _Platyzosteria ingens_ (on seventh segment).

_e._ _At the end of the body._

_Colænis julia_ ♀ (F. Müller), _Heliconius apseudes_ (F. Müller).

LITERATURE ON DEFENSIVE OR REPUGNATORIAL GLANDS

=Aldrovandus, U.= De animalibus insectis libri septem cum singulorum
iconibus ad vivum expressis. (Denuo impress: Bonon. apud Clementem
Ferronium, 1638, p. 273. The first edition was in 1602.)

=Moufet, T.= Insectorum sive minimorum animalium theatrum, etc.
London, 1634, pp. 185, 186.

=Gœdart, J.= Metamorphosis naturalis sive insectorum historia, etc.
Amstelodami, 1700, Pars ii, p. 136. (French ed. of 1700, ii, p. 162;
Lister’s Latin ed., London, 1685, p. 60.)

=Réaumur, R. A. F.= Mémoires pour servir à l’histoire des insectes,
etc. Paris, 1736, ii, pp. 266–269, Pls. 21, 22. [ii, partie ii, pp.
21–23, of the Amsterdam ed. of 1737–1748.]

=De Geer, C.= Observation sur la propriété singulière qu’ont les
grandes chenilles à quatorze jambes et à double queue, du saule, de
seringuer de la liqueur. (Mém. sav. étrang, Paris, 1750, i, pp. 530,
531, Pl.; Goetze und Bonnet, etc., Auserlesene Abhandlungen, 1774,
p. 220.)

=Schaeffer, J. C.= Neuendeckte Theile an Raupen und Zweyfaltern, etc.
Regensburg, 1754.

=Sulzer, J. H.= Die Kennzeichen der Insekten, etc. Zürich, 1761, pp.
65–67, Taf. 5, Fig. 34.

=Müller, O. F.= Pile-larven med dobbelt Hale, og dens Phalæne, etc.
Kjöbenhavn, 1772, pp. 53–56, Pl. 2, Figs. 3–5.

=Bonnet, C.= Mémoire sur une nouvelle partie commune à plusieurs
espèces de chenilles. (Mém. math. d. savants étrang., Paris, 1755,
ii, pp. 44–52; Collection complète des œuvres de C. Bonnet, 1779,
ii, pp. 3–16.)

—— Mémoire sur la grande chenille à queue fourchue du saule, dans
lequel on prouve, que la liqueur que cette chenille fait jaillir,
est un véritable acide, et un acide très-actif. (Mém. math, de
savants étrang., Paris, 1755, ii, pp. 267–282; Collection complète
des œuvres de C. Bonnet, 1779, ii, pp. 17–24.)

=Amoreux, P. J.= Notice des insectes de la France, réputés venimeux,
etc. Paris, 1789, pp. 282–285.

=Schwarz, C.= Neuer Raupenkalender. Nürenberg, 1791, Abth. i, p. 59.

=Petzhold, C. P.= Lepidopterologische Beyträge. (L. G. Scriba’s
Beiträge zu der Insekten-geschichte, Frankfurt am Main, 1793, Heft
3, pp. 230–251.)

=Nouvelle Dictionnaire d’Hist. Nat.=, xv, p. 487. (Larva of
Hydrophilus ejects with a slight noise a fœtid and blackish fluid.)

=Rengger, Johann Rudolph.= Physiologische Untersuchungen über die
thierische Haushaltung der Insecten. Tübingen, 1817. (In the chapter
entitled Abgesonderte Säfte bei den Raupen, he speaks of the
glandular apparatus of the larva of _B. vinula_, noticing the
general form of the secretory sac, that it opens out in two muscular
evertible points, out of which the secretion is ejected.)

=Dufour, L.= Mémoire anatomique sur une nouvelle espèce d’insecte du
genre Brachinus. (Ann. de mus. d’histoire nat., xviii, 1811, pp.
70–81.)

—— Recherches anatomiques sur les carabiques et sur plusieurs autres
coléoptères. (Ann. d. Sci. Nat., 1826, viii, pp. 5–54.)

—— Mémoire sur les métamorphosis et l’anatomie de la _Pyrochroa
coccinea_. Glande odorifique. (Ibid., ii sér. Zoologie, xiii, 1840,
pp. 340, 341.)

—— Recherches anatomiques sur les Diptères. 1851, pp. 195, 313.
(Alimentary canal of Sepsis contains the seat of a “glande
odorifique.”)

=Kirby and Spence.= Introduction to entomology, etc. (2d ed., i, 1815;
London, 1818, ii, pp. 238, 239.)

=Lyonet, P.= Recherches sur l’anatomie et les metamorphoses de
différentes espèces d’insectes. Ouvrage posthume. Paris, 1832.

=Morren, C.= Mémoire sur l’émigration du puceron du pêcher (_Aphis
persicæ_), et sur les caractères et l’anatomie de cette espèce.
(Ann. Sci. Natur. Zool., 1836, vi, pp. 65–93, Pls. 6, 7.)

=Ratzeburg, J. T. C.= Die Forstinsekten, etc. (Theil i, Die Käfer,
etc., 1837, p. 246.)

=Aubé, C.= Note sur une sécrétion fétide d’_Eumolpus pretiosus_. (Ann.
Soc. Ent. Fr., 1837, i, vi; Bull., p. 58.)

=Lacordaire, J. S.= Introduction à l’entomologie. 1838, ii, p. 45.

=Meckel, von Hemsbach, Johann Friedrich.= Mikrographie einiger
Drüsenapparate der niederen Thiere. (Anat. Phys. u. wiss. Med.,
1846, pp. 1–73, Taf. 1–3; p. 46, Müller’s Archiv.)

=Stein, Friedrich.= Vergleichende Anatomie und Physiologie der
Insekten. Berlin, 1847.

=Leidy, Joseph.= History and anatomy of the hemipterous genus
Belostoma. (Journ. Acad. Nat. Sci. Philadelphia, Ser. 2, 1847, i,
Part i, pp. 57–67, Pl. 1.)

—— Odoriferous glands of invertebrata. (Proc. Acad. Philadelphia,
1849, iv, 234–236, 1 Pl.; Ann. and Mag. N. H., Ser. 2, 1850, v, pp.
154–156.)

=Chapuis et Candèze.= Catalogue des larves des coléoptères, etc. (Mém.
Soc. Sci. de Liège, 1853, viii, pp. 351–653, Pls. 1–9, pp. 611,
612.)

=Siebold, Carl Theodor.= Lehrbuch der vergleichenden Anatomie der
wirbellosen Thiere, 1848. (Burnett’s transl., Boston, 1854.)

=Burnett, Waldo Irving.= Translation of Siebold’s Anatomy of the
Invertebrates, 1854. (Note on the osmeteria of _Papilio asterias_,
which he regards as an odoriferous and defensive, rather than
tactile, organ, p. 415.)

=Karsten, H.= Bemerkungen über einige shaarfe und brennende
Absonderungen verschiedener Raupen. (Müller’s Archiv für Anat. Phys.
u. wiss. Med., 1848, pp. 375–382, Taf. 11, 12.) Describes the
poison-glands at the base of the spines of Saturnia larvæ.

—— Harnorgane des _Brachinus complanatus_. (Müller’s Archiv, 1848, pp.
367–376, Taf.)

=Laboulbéne, Alexandre.= Note sur les caroncules thoraciques du
Malachius. (Annales de la Soc. Ent. de France, 3^e Sér., vi, 1858,
pp. 521–528.)

=Saussure, Henri de.= Recherches zoologiques de l’Amerique centrale et
du Mexique. (6^e Partie, Études sur les Myriopodes et les Insectes,
Paris, 1860.)

=Gerstaecker, C. E. A.= Ueber das vorkommen von ausstülpbaren
Hautanhängen am Hinterleibe an Schaben. (Archiv f. Naturgesch.,
1861, xxi, pp. 107–115.)

=Liegel, Hermann.= Ueber den Ausstülpungsapparat von Malachius und
verwandten Formen. Inaug. Diss., Göttingen, pp. 31, 1 Taf. (n. d.,
since 1858 and before 1878.)

=Leydig, F.= Zur Anatomie der Insecten. (Archiv f. Anat. Phys. u.
wiss. Med., 1859, pp. 33–89, 149–183, Taf. 2–4, pp. 35 and 38.)

—— Ueber bombardier Käfer. (Biolog. Centralbl., x, 1890, pp. 395,
396.)

=Claus, C.= Ueber die Seitendrüsen der Larve von _Chrysomela populi_.
(Zeits. f. wissens. Zool., xi, 1861, pp. 309–314, Taf. xxv.)

—— Ueber Schutzwassen der Raupen des Gabelschwanzes. (Würzburger
Naturw. Zeitschrift, 1862, iii, xiv; Sitz. am., 28 Juni, 1862.)

=Rogenhofer, Alois.= Drei Schmetterlingsmetamorphosen. (Verhandlungen
der k. k. zoolog.-bot. Gesellschaft, Wien, xiii, 1862, pp.
1224–1230.)

=Fitch, Asa.= Eighth report on the noxious and other insects of ...
New York. (Trans. N. Y. State Agric. Soc., 1862, xxii, pp. 657–684),
p. 677. (Separate.)

=Guenée, Achille.= D’une organe particulier que présente une chenille
de Lycæna. (Annales Soc. Ent. de France, Sér. 4, 1867, pp. 665–668,
Pl. 13.)

=Landois, L.= Anatomie der Bettwanze, _Cimex lectularius_, mit
berücksichtigung verwandter Hemipterengeschlechter. (Zeitsch. f.
wissens. Zool., 1868, xvii, pp. 206–224, 218–223, Taf. 11, 12.)

=Studer, Theodor.= Mittheilungen der naturforsch. Gesellschaft in
Bern, 1872–1873, No. 792–811, p. 101.

=Candèze, E.= Les moyens d’attaque et de défense chez les insectes.
(Bull. Acad. royale de Belgique, 2 Sér., xxxviii, 1874, pp.
787–816.)

=Mayer, Paul.= Anatomie von _Pyrrhocoris apterus_. (Reichert und du
Bois-Reymond’s Archiv f. Anat. Phys., etc., 1874, pp. 313–347, 3
Taf.)

=Scudder, Samuel Hubbard.= Odoriferous glands in Phasmidæ. (Psyche, i,
pp. 137–140, Jan. 14, 1876; Amer. Nat., x, p. 256, April, 1876.)

—— Prothoracic tubercles in butterfly caterpillars. (Psyche, i, pp.
64, 168, 1876.)

—— Organs found near the anus of the ♀ pupa of Danais, which recall
the odoriferous organs mentioned by Burnett, transl. Siebold’s Comp.
Anat. as occurring in Argynnis and other genera. (Psyche, iii, p.
278, 1882, p. 453, note 22.)

—— Glands and extensile organs of larvæ of blue butterflies. (Proc.
Bos. Soc. Nat. Hist., xxxiii, pp. 357, 358, 1888.)

—— Butterflies of Eastern United States. i-iii, 1889.

—— New light on the formation of the abdominal pouch in Parnassius.
(Trans. Ent. Soc. London for 1892, January, 1893, pp. 249–253.)

=Müller, Fritz.= Die Stinkkölbchen der weiblichen Maracujáfalter.
(Zeitschr. f. wissens. Zool., 1877, xxx, pp. 167–170, Taf. 9.)

=Plateau, Félix.= Note sur une sécrétion propre aux coléoptères
dytiscides. (Ann. Soc. Ent. Belg., 1876, v, xix, pp. 1–10.)

=Edwards, William H.= Notes on _Lycæna pseudargiolus_ and its larval
history. (Can. Ent., x, Jan., 1878, pp. 1–14. Fig.)

—— On the larvæ of _Lycæna pseudargiolus_ and attendant ants. (Can.
Ent., x, July, 1878, pp. 131–136.)

—— Butterflies of North America, i-iv. Many Pls. Phil., 1868—.

=Voges, Ernst.= Beiträge zur Kenntniss der Juliden. (Zeitsch. f.
wissens. Zoologie, xxxi, p. 127, 1878. The scent-glands are
retort-shaped bodies, the necks of which open into _foramina
repugnatoria_.)

=Rye, E. C.= Secretion of water-beetles. (Ent. Month. Mag., xiv,
1877–1878, pp. 232, 233.)

=Forel, A.= Der Giftapparat und die anal Drüsen der Ameisen. (Zeits.
f. wissens. Zool., 1878, xxx, Suppl., pp. 28–68, Taf. 3, 4.)

=Saunders, William.= Notes on the larva of _Lycæna scudderi_. (Can.
Ent., x, Jan., 1878, p. 14.)

=Weismann, A.= Ueber Duftschuppen. (Zool. Anzeiger, 26th Aug., 1878,
Jahrg., i, pp. 98, 99.)

=Gissler, Carl Friedrich.= On the repugnatorial glands in Eleodes.
(Psyche, ii, Feb., 1879, pp. 209, 210.)

—— Odoriferous glands on the 5th abdominal segment in nymph of
_Lachnus strobi_. (Fig. 273, p. 804, of Packard’s Report on Forest
and Shade Tree Insects, 1890.)

=Brunner von Wattenwyl, K.= Ueber ein neues Organ bei den Acridiodeen.
(Verhandl. k. k. Zool. Bot. Gesells. Wien., 1879, xxix;
Sitzungsber., pp. 26, 27.)

—— Verhandl. k. k. Zool. Bot. Gesells. Wien. (A peculiar organ on hind
femora of Acridiidæ.)

=Rougement, P.= Observations sur l’organe détonnant du _Brachinus
crepitans_ Oliv. (Bull. Soc. Sci. Nat. Neuchâtel, 1879, xi, pp.
471–478, Pl.)

=Goossens, Th.= Sur une organe entre la tête et la première paire de
pattes de quelques chenilles. (Ann. Soc. Ent. France, ix, p. 4,
1809; Bull., pp. 60, 61.)

—— Des chenilles vésicantes. (Ann. Ent. Soc. France, vi, pp. 461–464,
1887.)

=Coquillett, D. W.= On the early stages of some moths. (Can. Ent.,
March, 1880, xii, pp. 43–46.)

=Chambers, Victor Tousey.= Notes upon some Tineid larvæ. (Psyche, iii,
July, 1880, p. 67. Certain retractile processes “from the sides of
certain segments of the larva.”)

—— Further notes on some Tineid larvæ. (Psyche, iii, p. 135, Feb. 12,
1881. Larva of Phyllocnistis has eight pairs of lateral pseudopodia
on first eight abdominal segments.)

=French, G. H.= Larvæ of _Cerura occidentalis_ Lint, and _C. borealis_
Bd. (Can. Ent., July, 1881, xiii, pp. 144, 145.)

=Passerini, N.= Sopra i due tubercoli abdominali della larva della
_Porthesia chrysorrhœa_. (Bull. Soc. Ent. Ital., 1881, xiii, pp.
293–296.)

=Klemensiewicz, Stanislaus.= Zur näheren Kenntniss der Hautdrüsen bei
den Raupen und bei _Malachius_. (Verhandlungen d. Zool. Bot.
Gesellsch. Wien., xxxii, pp. 459–474, 1882, 2 Taf.)

=Weber, Max.= Ueber eine Cyanwasserstoffsäure bereitende Drüse.
(Archiv für Mikr. Anat., xxi, pp. 468–475, xxiv, 1882.)

=Bertkau, Philip.= Ueber den Stinkapparat von _Lacon murinus_ L.
(Archiv f. Naturg., 1882, Jahrg., xlviii, pp. 371–373.)

=Dimmock, George.= Organs, probably defensive in function, in the
larva of _Hyperchiria varia_ Walk. (_Saturnia io_ Harris). (Psyche,
iii, pp. 352, 353, Aug. 19, 1882. Account of lateral eversible
glands on 1st and 7th abdominal segments; they emit neither moisture
nor odor.)

—— On some glands which open externally on insects. (Psyche, iii, pp.
387–399, Jan. 15, 1883. Treats of poison-glands, glandular hairs,
eversible glands of Cerura, etc.)

=Coleman, N.= Notes on _Orgyia leucostigma_. (Papilio,
November-December, 1882, Jan., 1883, ii, pp. 164–166.)

=Müller, F.= Der Anhang am Hinterleibe der _Acræa_-weibchen. (Zool.
Anzeiger, 6th Aug., 1883, Jahrg., vi, pp. 415, 416.)

=Dewitz, H.= Ueber das durch die Foramina repugnatoria entleerte
Secret bei Glomeris. (Biol. Centralblatt, iv, pp. 202, 203, 1884.)

=Williston, S. A.= Protective secretion of Eleodes ejected from anal
gland. (Psyche, iv, p. 168, May, 1884.)

=Poulton, Edward Bagnall.= Notes in 1885 upon lepidopterous larvæ and
pupæ, including an account of the loss of weight in the
freshly-formed lepidopterous pupæ. (Trans. Ent. Soc., London, June,
1886, pp. 156, 157, 159.)

—— Notes in 1886 upon lepidopterous larvæ, etc. (Trans. Ent. Soc.,
London, Sept., 1887, pp. 295–301.)

—— Notes in 1887 upon lepidopterous larvæ, etc. (Trans. Ent. Soc.,
London, 1888, p. 597.)

=Künckel-d’Herculais, J.= La punaise de lit et ses appareils
odoriférants. (Comptes rendus, ciii, 1886, pp. 81–83; Annals & Mag.
Nat. Hist., 5th Ser., xviii, 1886, pp. 167, 168.)

—— Étude comparée des appareils odorifiques dans les differents
groupes d’Hemiptères hétéroptères. (Compt. rend. Acad. Sc., Paris,
cxx, pp. 1002–1004.)

=Packard, A. S.= The fluid ejected by notodontian caterpillars. (Amer.
Nat., 1886, xx, pp. 811, 812.)

—— An eversible “gland” in the larva of Orgyia. (Amer. Nat., 1886, xx,
p. 814.)

—— Fifth Rep. U. S. Ent. Comm. Insects injurious to forest and shade
trees, p. 136, 1890.

—— Hints on the evolution of the bristles, spines, and tubercles of
certain caterpillars. (Proc. Boston Soc. Nat. Hist., xxiv, 1890, p.
551.)

—— Notes on some points in the external structure and phylogeny of
lepidopterous larvæ. (Proc. Bost. Soc. Nat. Hist., xxv, 1890, pp.
83–114.)

—— A study of the transformations and anatomy of _Lagoa crispata_, a
bombycine moth. (Proc. Amer. Phil. Soc., Philadelphia, xxxii, 1893,
pp. 275–292, 7 Pls.)

—— The eversible repugnatorial scent glands of insects. (Journ., N. Y.
Ent. Soc. iii, 1895, pp. 110–127; iv, p. 896; pp. 26–32, 1 Pl.)

=Loman, J. C. C.= Freies Jod als Drüsensecret. (Tijdschr. Neder.
Dierk. Ver. Deel 1, 1887, pp. 106–108.)

=Riley, Charles Valentine.= Proc. Ent. Soc., Washington, March 13,
1888, i, pp. 87–89.

—— Notes on the eversible glands of larvæ of _Orgyia_ and _Parorgyia
leucopæa_ and _P. clintonii (achatina)_. (See 5th Rep. U. S. Ent.
Comm., p. 137.)

=Denham, Ch. S.= The acid secretion of _Notodonta concinna_. (Insect
Life, i, p. 147, 1888; hydrochloric acid.)

=Michin, Edward A.= Note on a new organ, and on the structure of the
hypodermis, in _Periplaneta orientalis_. (Quart. Journ. Micros. Sc.,
Dec., 1888, xxiv, 1 Pl.)

—— Further observations on the dorsal gland in the abdomen of
Periplaneta and its allies. (Zool. Anz., 27 Jan., 1890, pp. 41–44.)

=Maynard, C. L.= The defensive glands of a species of Phasma,
_Anisomorpha buprestoides_. (Contributions to Science, i, April,
1889.)

=Schaeffer, Cæsar.= Beiträge zur Histologie der Insekten. (Zool.
Jahrb. Morph. Abth. iii, pp. 611–652, Taf. xxix, xxx, 1889; treats
of the ventral glands in prothorax of caterpillars; scales and hairs
are secretions from the very greatly enlarged hypodermic cells.)

=Gilson, G.= Les glandes odorifères der _Blaps mortisaga_ et de
quelques autres espèces. (La Cellule, v. pp. 1–21, 1 Pl., 1889.)

—— The odoriferous apparatus of _Blaps mortisaga_. (Rep. 58th Meeting
Brit. Assoc. Adv. Sc., 1889, pp. 727, 728.)

=Haase, Erich.= Ueber die Stinkdrüsen der Orthoptera. (Sitzgsber. Ges.
Naturf. Freunde, Berlin, pp. 57, 58, 1889.)

—— Zur Anatomie der Blattiden. (Zool. Anz., xii Jahrg., pp. 169–172,
1889.)

=Herbst, Curt.= Anatomische Untersuchungen an _Scutigera coleoptrata_.
Ein Beitrag zur vergleichenden Anatomie der Articulaten. Dissert.,
Jena, pp. 36 (Hautdrüsen, Coxal-Organ.); p. 1, 1889.

=Wheeler, William M.= Hydrocyanic acid secreted by _Polydesmus
virginiensis_ Drury. (Psyche, v, p. 422.)

—— New glands in the hemipterous embryo. (Amer. Nat., Feb. 1890, p.
187; odorous(?) glands.)

=Jackson, W. Hatchett.= Studies in the morphology of the Lepidoptera,
Pt. i. (Trans. Linn. Soc., London, 2 Ser., Zoöl., v, May, 1890.)

=Krauss, Hermann.= Die Duftdrüse der _Aphlebia bivittata_ Brullé
(Blattidæ) von Teneriffa. (Zool. Anz., xiii Jahrg., 1890, pp.
584–587, 3 Figs.)

=Fernald, H. T.= Rectal glands in Coleoptera. (Amer. Nat., xxiv, pp.
100, 101, Pls. 4, 5, 1890.)

=Verson, E.= Hautdrüsen system bei Bombyciden (Seidenspinner). (Zool.
Anzeiger, 1890, pp. 118–120.)

=Vosseler, Julius.= Die Stinkdrüsen der Forficuliden. (Arch. Mikr.
Anat., xxxvi, 1890, pp. 565–578, Taf. 29.)

=Carrière, J.= Die Drüsen am ersten Hinterleibsringe der
Insektenembryonen. (Biol. Centralblatt, xi, pp. 110–127, 1891.)

=Borgert, Henry.= Die Hautdrüsen der Tracheaten. Inaugural Diss.,
Jena, 1891, pp. 1–80.

=Lang, Arnold.= Lehrbuch der vergleichende Anatomie, English Trans. by
Henry M. and Matilda Bernard, 1891, pp. 458, 459.

=Kennel, J. von.= Die Verwandtschaftverhältnisse der Arthropoden.
(Schriften herausgegeben von der Naturforscher Gesellschaft bei der
Universität Dorpat, vi, Dorpat, 1891.)

=Patton, W. H.= Scent-glands in the larva of Limacodes. (Can. Ent.,
xxiii, Feb. 1891, pp. 42, 43; eight pairs of glands with pores along
the edges of the back.)

=Batelli, Andrea.= Di una particolarita nell integumento dell’
_Aphrophora spumaria_. (Monitore Zoöl. Ital. Anno 2, pp. 30–32,
1891. Dermal glands in the hindermost segment.)

=Ash, C. D.= Notes on the larva of _Danima banksii_ Lewin. (Ent.
Month. Mag., Sept. 1892, p. 232, Fig.) notodontian larva protrudes
from under side of prothoracic segment a Y-shaped, red organ like
that of Papilio; no odor or fluid given out.

=Bernard, Henry M.= An endeavor to show that the tracheæ of the
Arthropoda arose from setiparous sacs. (Spengel’s Zool., Jahrbuch,
1892, pp. 511–524, 3 Figs.)

=Latter, Oswald.= The secretion of potassium hydroxide by _Dicranura
vinula_, and the emergence of the imago from the cocoon. (Trans.
Ent. Soc. London, 1892, 287, also xxxii; Prof. Meldola adds that the
larva of _D. vinula_ secretes strong, formic acid, and is the only
animal known to secrete a strong, caustic alkali.)

—— Further notes on the secretion of potassium hydroxide by _Dicranura
vinula_ (imago), and similar phenomena in other Lepidoptera. (Trans.
Ent. Soc. London; Nature, 1895, p. 551, March 20, 1895.)

=Zograff, Nicolas.= Note sur l’origine et les parentes des
Arthropodes, principalement des Arthropodes trachèates. (Congrès
Internationale de Zoologie, 2^e Session à Moscow, Aug. 1892; Part i,
Moscow, 1892, pp. 278–302, 1892; cyanogenic glands in Myriopods, p.
287.)

=Swale, H.= Odor of _Olophrum piceum_. (Ent. Month. Mag., v, Jan.
1896, pp. 1, 2.)

=Cuénot, L.= Moyens de défense dans la série animale, Paris, n. d.
(1892); the ejection of blood as a means of defence by some
Coleoptera. (Comptes rendus, Acad. Sc. France, April 16; Nature,
April 26, 1894.)

—— Sur la saignée réflexe et les moyens de défense de quelques
insectes. (Arch. Zool. expér. (3), 1897, iv, pp. 655, 679, 680.)

=Holmgren, Emil.= Studier öfverhudens och de körtelartade hudorganens
morfologi hos skandinaviska macrolepidopterlarver. (K. Svenska
Vetenskaps-Akademiens Handlingar, xxvii, No. 4, Stockholm, 4º 1895,
pp. 82, 9 Pls.)

=Lutz, K. G.= Das Blut der Coccinelliden. (Zool. Anzeiger, 1895, pp.
244–255, 1 Fig.)

=Gilson, Gustav.= Studies in insect morphology. (Proc. Linn. Soc.
London, March 5, 1896; Nature, p. 500.)

—— On segmentally disposed thoracic glands in the larvæ of the
Trichoptera. (Journ. Linn. Soc., London, xxv. 1897.)

=Cholodkowsky, N.= Entomotomische Miscellen, v, Ueber die
Spritzapparate der Cimbiciden Larven, pp. 135–143, 2 Taf. Ibid., vi.
Ueber das Bluten der Cimbiciden Larven, pp. 352–357, 1 Fig. (Horæ
Soc. Ent. Rossicæ, xxx, 1897.)

Also the writings of Darwin, Wallace, Poulton, Weir, Beddard, Butler,
Busgen, (pp. 365–367), Girard, Kolbe, Locy.

THE ALLURING OR SCENT-GLANDS

It is difficult to draw the line between repelling and alluring glands.
Attention was first definitely called to the alluring odors of
Lepidoptera by Fritz Müller, who showed that the males of certain
butterflies are rendered attractive to the other sex by secreting
odorous oils of the ether series. He pointed out that the seat of the
odor is the androconia (see p. 199), while either repellent or pleasant
odors are exhaled from abdominal glands.

Those of _Pieris napi_ yield a scent like that of citrons, _Didonis
biblis_ gives off three different odors from different parts of the
body, besides having a distinctly odorous spot on the hind wings.
Both sexes have a sac between the fourth and fifth abdominal
segments which exhales a very unpleasant (protective) odor, while
the males have on the succeeding segment a pair of glands from which
proceeds an agreeable odor like that of the heliotrope. _Callidryas
argante_ throws off a musky odor. In _Prepona laertes_ the odor is
like that of a bat, in _Dircenna xantho_ it is vanilla-like, the
androconia being situated on the front edge of the hind wings. In
_Papilio grayi_ the odor is said to be as agreeable and intense as
in flowers. Certain sphingids are known to exhale a distinct odor,
which Müller has traced to a tuft of hair-like scales at the base of
the abdomen, and which fits into a groove in the first segment, so
as to be ordinarily invisible.

[Illustration:

_Fig. 367._—Scent-tufts: 1, of _Leucarctia acræa_; 2, of _Pyrrarctia
isabella_.—After Smith.
]

In the noctuid genus, Patula, the costal half of the hind wing is
modified to form a large scent-gland, and in consequence the
venation has been modified. The still greater distortion of the
veins in the allied genus, Argida, was attributed by the author to
its once having possessed a similar scent-gland, now become
rudimentary by disuse. (Hampson.)

Peculiar white or orange-colored, hairy, thread-like processes have
been found protruding from narrow openings near the tip of the
abdomen of Arctian moths (Fig. 367), which throw off, according to
J. B. Smith, “an intense odor, somewhat like the smell of laudanum.”
We have perceived the same unpleasant odor emanating from the males
of _Spilosoma virginica_ and _Arctia virgo_, as well as _Leucarctia
acræa_.

We are informed by C. Dury that similar but longer hairy appendages
are thrust out by the male of _Haploa clymene_. Many glaucopid moths
protrude similar glandular processes. Thus Müller tells us that on
seizing a glaucopid female by the wings, nearly the whole body
became enveloped in a large cloud of snow-white wool which came out
of a sort of pouch on the ventral side of the abdomen.

The male of a glaucopid was seen to dart out a pair of long hollow
hairy retractile filaments which in some species exceed the whole
body in length. The apparatus secretes a peculiar odor, probably
serving to allure the female (Nature), and certain Zygænidæ have on
the inner side of the paranal lobes (Afterklappen) glands filled
with a sweetly scented fluid. Smith has detected a peculiar brush of
hair-like scales in a groove between the dorsal and ventral parts of
the basal two segments of the abdomen of _Schinia marginata_ (family
Noctuidæ), and when removed it exhaled a laudanum-like smell.

The pupa of _Citheronia regalis_ gives out from the end of the
abdomen a scent reminding us of laudanum.

[Illustration:

FIG. 368.—Scent-tufts on middle legs of _Catocala concumbens_.—After
Bailey.
]

Another mode of disseminating pleasant, alluring odors is that of the
males of certain moths, which bear pencils and tufts on their fore or
hind legs, and in the case of an Indian butterfly on the greatly
elongated palpi. Those on the legs are ordinarily concealed in cavities
or furrows in the leg, and may be thrust out and expanded so as to
widely diffuse their odor. Such are those of the males of Catocala (Fig.
368), which resemble an artist’s fitch brush. In _Hepialus hecta_, where
the arrangements for protecting the tufts are quite abnormal, Bertkau
has detected the cells which secrete the odorous fluid. In the male of
another Hepialus (_H. humuli_) a peculiar scent proceeds from the
curiously aborted and altered hind tibiæ. (Barrett.) In one case, that
of a geometrid moth (_Bapata dichroa_ of New Guinea), these pencils
occur on all the legs. (Haase). In many species a distinct odor is
perceptible when the leg bearing the pencil or tuft is crushed.

These eversible scent-glands have been supposed to be mostly restricted
to the Lepidoptera, and to a single known case in the Trichoptera, but
similar alluring male glands also occur in the Orthoptera (Locustidæ).
H. Garman has described and figured in the cave cricket (_Hadenœcus
subterraneus_) “a pair of white fleshy appendages protruding from slits
between the terga of the 9th and 10th abdominal somites, the nature of
which is not clear,” adding, “the slits through which the organs appear
are situated one on each side anterior to and a little within the cerci.
When fully protruded, the glands are white, cylindrical, a little
tapering, and are about one-eighth of an inch long.” He believes that
they are protruded during the period of sexual excitement, and suggests
that “the sense of smell is certainly the one best calculated to bring
the sexes together in the darkness of caves.” We had previously noticed
these organs in alcoholic specimens, but supposed that they were fungous
growths. On dissecting and making microscopic sections of them, the
gland is, when extended (Fig. 369), seen to be a long, ensiform, sharp,
band-like process, with numerous retractor muscular fibres. When at rest
each gland is folded about five times, forming a bundle lying on each
side of the end of the intestine. The walls are formed of a single layer
of epithelium, as seen in Fig. 369, _B_.

[Illustration:

FIG. 369.—Eversible scent-glands (_a_) of Hadenœcus, nat. size:
Kingsley, _del._; _A_, a gland outstretched, with the retractor
muscular fibres; _t_, part of the tergite. _B_, section of the
gland, showing the single layer of epithelial cells, and the
muscular fibres (_m_).—Author _del._
]

In the male of the common wingless cricket, _Ceuthophilus maculatus_, we
have discovered what appears to be a pair of scent-glands lying directly
over the last abdominal ganglion. They form two large white sacs
situated close together, with a short common duct which passes back and
opens externally upwards by a transverse slit on the under side of the
last segment of the body.

LITERATURE ON ALLURING GLANDS

=Watson, J.= On the microscopical examination of plumules, etc. (Ent.
Month. Mag., ii, 1865, p. 1.)

—— On certain scales of some diurnal Lepidoptera. (Mem. Lit. and Phil.
Soc. Manchester, Ser. 3, ii, 1868, p. 63.)

—— On the plumules or battledore scales of Lycænidæ. (Mem. Lit. and
Phil. Soc. Manchester, Ser. 3, iii, 1869, p. 128.) Further remarks,
etc. (Ibid., p. 259.)

=Anthony, J.= Structure of battledore scales. (Month. Microsc. Journ.,
vii, 1872, p. 250; see also p. 200.)

=Morrison, Herbert Knowles.= On an appendage of the male _Leucarctia
acræa_. (Psyche, i, pp. 21–22, October, 1874.)

=Müller, Fritz.= The habits of various insects. (Nature, June 11,
1874, pp. 102–103.)

—— Ueber Haarpinsel, Fitzflecke und ähnliche Gebilde auf den Flügeln
männlicher Schmetterlinge. (Jena. Zeitschr. f. Naturw., 1877, xi,
pp. 99–114.)

—— Beobachtungen an brasilianischen Schmetterlingen, ii. I. Die
Duftschuppen der männlichen Maracujáfalter. (Kosmos, 1877, i, pp.
391–395, Figs. 5, 6.) II. Die Duftschuppen des männchens von _Dione
vanillæ_. (Kosmos, ii, 1877, pp. 38–42, 7 Taf.)

—— As maculas sexuaes dos individuos masculinos das especies _Danais
erippus_ e _D. gilippus_. (Arch. Mus. Nac. Rio Janeiro, ii, 1877
(1878), pp. 25–29, 1 Pl.)

—— Die Duftschuppen der Schmetterlinge (nach dem “Kosmos” in Ent.
Nachr., 1878, pp. 29–32, 109).

—— Wo hat der Moschusduft der Schwärmer seinen Sitz? (Kosmos, ii
Jahrg., 1878, pp. 84, 85.)

—— Os orgaos odoriferos dos especias _Epicalia acontius_, Lin. e de
_Myscelia orsis_, Dru. (Arch. Mus. Nac. Rio Janeiro, ii, 1879, pp.
31–35.)

—— Os orgaos odoriferos nas pernas de certos Lepidopteres. (Arch. Mus.
Nac. Rio Janeiro, ii, 1879, pp. 37–46, 3 Pls.)

—— Os orgaos odoriferos da _Antirrhœa archœa_. (Arch. Mus. Nac. Rio
Janeiro, iii, 1878, pp. 1–7, 1 Pl.)

—— A prega costal das Hesperideas. (Arch. Mus. Nac. Rio Janeiro, iii,
1880, pp. 41–50, 2 Pls.)

=Weismann, August.= Ueber Duftschuppen. (Zool. Anzeiger, i, 1878, pp.
98, 99.)

=Arnhart, L.= Sexundäre Geschlechtscharaktere von _Acherontia
atropos_. (Verh. d. k. k. zool. bot. Ges. Wien, xxix, 1879, p. 54.)

=Bertkau, Philipp.= Duftapparat an Schmetterlingsbeinen. (Ent.
Nachrichten, 1879, Jahrg., pp. 223, 224.)

—— Ueber den Duftapparat von _Hepialus hecta_. (Archiv f. naturg.,
xlviii Jahrg., 1882, pp. 363–370, Figs.; also in Biol.
Centralblatt., ii Jahrg., 1882, pp. 500–502.)

—— Ergänzung (Duftvorrichtungen bei Lepidopteren). (Ent. Nachr., 1880,
p. 206.)

—— Entomologische Mizellen. 1. Ueber Duftvorrichtungen einiger
Schmetterlinge. (Verh. d. naturhist. Ver. d. preuss. Rheinlande und
Westf., 1884, pp. 343–350.)

=Reichenau, W. von.= Der Duftapparat von _Sphinx ligustri_. (Ent.
Nachr., 1880, p. 141; also Kosmos, iv Jahrg., 1880, pp. 387–390.)

=Fügner, R.= Duftapparat bei _Sphinx ligustri_. (Ent. Nachr., 1880, p.
166.)

=Lelievre, Ernest.= (Note in Le Naturaliste, June 1, 1880. Both sexes
of _Thais polyxena_ emit an odorous exhalation. Notes on exhalation
from _Spilosoma fuliginosa._)

=Hall, C. G.= Peculiar odor emitted by _Acherontia atropos_.
(Entomologist, London, xvi, p. 14.)

=Åurivillius, Christopher.= Ueber secundäre Geschlechtscharactere
nordischer Tagfalter. (Stockholm, 1880, Bihang till K. Svensk. Vet.
Akad. Handl., v, pp. 56, 3 Taf.)

—— Des caractères sexuels secundaires chez les papillons diurnes.
(Ent. Tidskrift, 1880, pp. 163–166.)

—— Anteckningar om några skandinaviska fjärilarter. (Ent. Tidskr., iv,
Årg., 1884, pp. 33–37; Résumé (French), ibid., pp. 55–57.)

=Kirby, W. F.= Fans on the fore legs of _Catocala fraxini_. (Papilio,
ii, p. 84, 1882.)

=Bailey, James S.= Femoral tufts or pencils of hair in certain
Catocalæ. (Papilio, ii, 1882, pp. 51, 52, 146; also in Stettin Ent.
Zeitung, xliii, p. 392.)

=Edwards, Henry.= Fans on the feet of Catocaline moths. (Papilio, ii,
p. 146, 1882.)

=Stretch, R. H.= Anal appendages of _Leucarctia acræa_. (Papilio, iii,
pp. 41, 42, 1883, 1 Fig.)

=Weed, Clarence M.= Appendages of Leucarctia. (Papilio, iii, 1883, p.
84.)

=Grote, Aug. R.= Appendages of _Leucarctia acræa_. (Papilio, iii,
1883, p. 84.)

=Haase, Erich.= Ueber sexuelle Charactere bei Schmetterlingen.
(Zeitschr. f. Ent., Breslau, N. F., 1885, pp. 15–19, 36–44; also
Ent. Nachr., xi Jahrg., pp. 332, 333.)

—— Duftapparate indo-australischer Schmetterlinge. (Corresp. Blatt.
Ent. Ver. Iris, Dresden, 1886, pp. 92–107, 1 Taf.; ibid., 1887, pp.
159–178; ibid., 1888, pp. 281–336.)

—— Ueber Duftapparate bei Schmetterlingen. (Sitzgsber. Nat. Ges. Iris,
Dresden, 1886, pp. 9–10; Abstr. in Journ. R. Micr. Soc., vi, pp.
969–970, 1886.)

—— Der Duftapparate von Acherontia. (Zeitschr. f. Ent., Breslau, N.
F., 1887, pp. 5–6.)

—— Dufteinrichtung indischer Schmetterlinge. (Zool. Anzeiger, 1888,
pp. 475–481.)

=Dalla Torre, K. W. von.= Die Duftapparate der Schmetterlinge.
(Kosmos, 1885, ii, pp. 354–364, 410–423; Abstr. by J. B. Smith in
Proc. Ent. Soc., Washington, i, pp. 38, 1888.)

=Smith, John B.= _Cosmosoma omphale._ (Entomologica Americana, i, pp.
181–185, 1886. Describes and figures cavities in under side of
2d–4th abdominal segments of male, filled with a silky substance.
This may be for display to attract ♀, as the whole mass must be very
conspicuous when protruded. No odor noticed.)

—— Scent organs in some Bombycid moths. (Entomologica Americana, ii,
No. 4, pp. 79–80, 1886. Describes and figures long, slender, forked
hairy, orange or white, eversible glands, everted from between 7th
and 8th segments of abdomen of ♂ of _Leucarctia acræa_, _Pyrrharctia
isabella_, _Scepsis fulvicollis_, and _Cosmosoma omphale_.)

—— [Notes on odors and odoriferous structures of various moths and a
note by L. O. Howard on odor of Dynastes.] (Proc. Ent. Soc.,
Washington, i, pp. 40, 55, 56.)

=Müller, W.= Duftorgane der Phryganiden. (Archiv f. Naturgesch., 1887,
Jahrg. liii, pp. 95–97.)

=Pollack, W.= Duftapparate der _Hadena atriplicis_ und Litargyria. (xv
Jahrb. Westphäl. Prov. Ver. Münster, 1887, p. 16.)

=Patton, W. H.= Scent-glands in the larva of Limacodes. (Can. Ent.,
1891, xxiii, pp. 42, 43.)

=Garman, H.= On a singular gland possessed by the male _Hadenœcus
subterraneus_. (Psyche, 1891, p. 105, 1 Fig.)

=Barrett, C. G.= Scent of the male _Hepialus humuli_. (Ent. Month.
Mag., Ser. 2, iii, 1892, p. 217. Arises from the curiously aborted
and altered hind tibiæ.)

Also the writings of Baillif, Duponchel, F. Müller, Scudder (Psyche,
iii, p. 278, 1881), Burgess, Keferstein, Alpheraky, Plateau,
Marshall and Nicéville, Wood-Mason, White, Hampson.

THE ORGANS OF CIRCULATION

Although Malpighi was the first to discover the heart in the young
silkworm, it was not until 1826 that Carus proved that there was a
circulation of blood in insects, which he saw flowing along each side of
the body, and coursing through the wings, antennæ, and legs of the
transparent larva of Ephemera, though three years earlier Herold
demonstrated that the dorsal vessel of an insect is a true heart,
pulsating and impelling a current of blood towards the head. This
discovery was extended by Straus-Dürckheim, who discovered the
contractile and valvular structures of the heart. It is noteworthy that
both Cuvier and Dufour denied that any circulation, except of air,
existed in insects; and so great an anatomist as Lyonet doubted whether
the dorsal vessel was a genuine heart, though he pointed out the fact
that there are no arteries and veins connected with this vessel. Another
French anatomist, Marcel de Serres, thought that the dorsal vessel was
merely the secreting organ of the fat-body.

The so-called peritracheal circulation claimed by Blanchard and by
Agassiz has been shown by McLeod to be an anatomical impossibility,
the view having first been refuted by Joly in 1849.

Except the aorta-like continuation in the thorax and head which
divides into two short branches, there are, with slight exceptions
(p. 405), no distinct arteries, such as are to be found in the
lobster and other Crustacea, and no great collective veins, such as
exist in Crustacea and in Limulus. This is probably the result of a
reduction by disuse in the circulatory system, since in myriopods
(Julidæ and Scolopendridæ) lateral arteries are said to diverge near
the ostia.

_a._ The heart

The heart or “dorsal vessel” is a delicate, pulsating tube, situated
just under the integument of the back, in the median line of the body,
and above the digestive canal. It can be partially seen without
dissection in caterpillars. It is covered externally and lined within by
membranes which are probably elastic; and between these two membranes
extends a system of delicate muscular fibres, which generally have a
circular course, but sometimes cross each other. The heart is divided by
constrictions into chambers, separated by valvular folds. The internal
lining membrane referred to forms the valvular folds separating the
chambers. Each of these chambers has, at the anterior end, on each side,
a valvular orifice (Fig. 370, ostium, _i_) which can be inwardly closed.

Miall and Denny thus describe the different layers of the wall of
the heart of the cockroach:

“There are: (1) a transparent, structureless intima, only visible
when thrown into folds; (2) a partial endocardium, of scattered,
nucleated cells, which passes into the interventricular valves; (3)
a muscular layer, consisting of close-set, annular, and distant,
longitudinal fibres. The annular muscles are slightly interrupted at
regular and frequent intervals, and are imperfectly joined along the
middle line above and below, so as to indicate (what has been
independently proved) that the heart arises as two half-tubes, which
afterwards join along the middle. Elongate nuclei are to be seen
here and there among the muscles. The adventitia (4), or connective
tissue layer, is but slightly developed in the adult cockroach.”

[Illustration:

FIG. 370.—Part of the heart of _Lucanus cervus_: _a_, the posterior
chambers (the anterior ones are covered by a part of the ligaments
which hold the heart in place); _i_, auriculo-ventricular openings;
_g_, _g_, the lateral muscles fixed by the prolongations _h_, _h_,
to the upper side of the abdomen.—After Straus-Dürckheim.
]

Graber says that the heart of insects may be regarded not as an organ
_de novo_, but only as the somewhat modified contractile dorsal vessel
of the annelids, in which, however, the transverse arteries arising on
each side became, with the gradual development of the tracheæ,
superfluous and finally abortive. He describes it as a muscular tube
composed of very delicate annular fibres, which within and without is
covered by a relatively homogeneous, strong, elastic membrane.

The division into separate chambers is effected by means of a
folding inwards and forwards of the entire muscular wall. “A portion
of each side of the heart is first extended inwards so as very
nearly to meet a corresponding portion from the opposite side, and
then, being reflected backwards, forms, according to Straus
(Consid., etc., p. 356), the interventricular valve which separates
each chamber from that which follows it. Posteriorly to this valve,
at the anterior part of each chamber, is a transverse opening or
slit (Fig. 371, _b_), the _auriculo-ventricular orifice_, through
which the blood passes into each chamber, and immediately behind it
is a second, but much smaller, _semilunar valve_ (_c_), which, like
the first, is directed forwards into the chamber. It is between
these two valves on each side that the blood passes into the heart,
and is prevented from returning by the closing of the semilunar
valve. When the blood is passing into the chamber, the
interventricular valve is thrown back against the side of the
cavity, but is closed when, by the contraction of the transverse
fibres, the diameter of each chamber is narrowed, and the blood is
forced along into the next chamber.” (Newport.)

[Illustration:

FIG. 371.—A, heart of _Lucanus cervus_: _a_, valves or chambers;
_bb_, alary muscles; _c_, supposed auricular space around the
heart. _B_, division into arteries of the end of the aorta in
larva of _Vanessa urticæ_. _C_, interior of the chamber, showing
the transverse fibres; _b_, auriculo-ventricular opening and valve
into the chambers; _c_, semilunar valve; _d_, interventricular
valve.—After Straus-Dürckheim, from Newport.
]

[Illustration:

FIG. 372.—Heart of Belostoma.—After Locy.
]

According to Müller, there is but a single pair of ostia in Phasma,
and, in the larva of Corethra, the heart is a simple, unjointed
tube, not divided into chambers, and Viallanes states that, in the
very young larva of Musca, there are no ostia (Kolbe). In the larva
of Ptychoptera, Grobben found a short oval heart, with one pair of
ostia situated in the 6th abdominal segment; a long aorta proceeds
from it, the thoracic portion of which pulsates; from behind the
heart arises a pulsating pouch, which connects with the hinder
aorta, which does not pulsate, and ends at the base of two tracheal
gills. Burmeister was able to find only four pairs of openings in
the larva of Calosoma. Newport states that, while Straus figures
nine chambers in Melolontha, and, consequently, eight pairs of
openings, he has not been able to observe more than seven pairs of
openings in _Lucanus cervus_. He has invariably found eight pairs of
openings both in the larva and imago of _Sphinx ligustri_, as well
as in other Lepidoptera. According to Béla-Dezso, the number of
pairs of ostia corresponds to that of the pairs of stigmata.

There also occur, on each side of each chamber, two so-called
pear-shaped bodies which are separated from the tubular portion of
the heart itself, but, by means of muscular fibres, are united with
the chamber and with their valves. These pyriform bodies appear as
vesicles or cells with granular contents, besides some nuclei with
nucleoli. They are of very small size. According to the measurements
of Dogiel, in the larva of _Corethra plumicornis_, they are 0.02 to
0.1 mm. long, and 0.06 to 0.08 mm. broad. He regards these peculiar
bodies as apolar nerve-cells of the heart. (Kolbe.)

[Illustration:

FIG. 373.—A, part of the heart of _Dyticus marginalis_, showing the
spiral arrangement of the muscular fibres; _c_, closed, _e_, open,
valve; _a_, dorsal diaphragm with interwoven muscular fibres; _b_,
arrangement of fibres, recalling the screw-like features of the
fibres of the human heart; _d_, narrow end. _B_, diagrammatic
figure of the valvular openings, with the terminal flap (_e_), and
the cellular valve, of a May beetle; _a_, valvular opening of a
dipterous larva, with the interventricular valve (_b_). _C_,
abdomen of a mole-cricket, ventral view; _c_, the segmented heart;
_a_, aorta; _b_, segmented diaphragm under it.—After Graber.
]

Besides the venous openings of the heart which open into the
pericardial region, Kowalevsky has discovered, in the heart of some
Orthoptera (Caloptenus, Locusta, etc.), five pairs of openings by
which the cardiac chambers receive the blood of the peri-intestinal
region. Graber had divided the cœlom of insects into three regions
(pericardial, peri-intestinal, and perineural regions), and hitherto
only a union of the heart with the pericardial region by slit-like
openings was known. These openings are symmetrically distributed on
five abdominal segments; each section of the heart in this region
has, therefore, four openings, which are all of a truly venous
nature. These openings, called cardio-cœlomic apertures, are visible
to the naked eye, being situated on conical papillæ of the walls of
the heart. These papillæ pass through the outer diaphragm, and open
into the peri-intestinal part of the cœlom, in the Acrydiidæ
directly, in the Locustidæ through special canals. The cells of the
papillæ are spongy, possessing large nuclei, and similar, as a
whole, to glandular cells. (Comptes rendus, cxix, 1894.)

The mechanism by which the ostia are closed consists, according to
Graber, of an ∞-shaped muscle passing around the two openings, and
which, being interlaced, is sufficient to close the openings. But
this is not all. The fore and hinder edge of the ostia project,
leaf-like, into the cavity of the heart, and thus form, with the
outer walls, two valves which, during the systole, filled with the
blood rushing in, not only hermetically close the lateral openings,
but also, by the simultaneous closure of the entire chamber by the
circular muscles in the middle of the same, the two valves,
simultaneously approaching each other, so nearly touch that they
form a transverse partition wall in the chamber. But, for the last
purpose, _i.e._ for the separation of the chambers from one another,
there is a very special contrivance. In the May beetle, we find,
besides a valve (Fig. 373, _B_, _e_), opening into the middle of the
chambers, a large, stalked cell (_d_), which, in the diastole,
_i.e._ in the expansion of the heart, hangs down free on the walls
of the heart; but, in the systole or contraction, like a cork,
closes the middle of the valve, but does not wholly close the
cavity. He has observed, in the larva of Corethra, formal,
interventricular valves, which also are not in the middle, but are
separated from one another in the interlaced ends. They consist of
two longitudinally membranous flaps which move against each other
like two valves (Fig. 373, _B_, _b_).

“But what is the necessity for such a complicated mechanism? All the
blood from behind passes into the heart, and, for its propulsion a
simple muscular tube, whose circular fibres would draw together and
contract it, would be thought to be sufficient. But the heart,
except in some larvæ, ends posteriorly in a blind sac, and the blood
can only pass into it by a series of pairs of lateral openings. Now,
as regards the reception and the propulsion of the blood forwards,
two modes are conceivable. The simplest way would be that the
tubular heart should, along its whole length, contract or expand;
that, moreover, the blood should be simultaneously sucked in through
all the openings, and that then, also, the contraction, or systole,
should take place in every part of the heart at the same moment. But
this would, plainly, in so long and thin-walled a vessel, be highly
impracticable, since, through such a manipulation, the mass of blood
enclosed in the heart would be crowded together rather than really
impelled forwards. Only the second case could be admissible, and
that is this, that each chamber pulsates, one after another, from
behind forwards. But, then, each segmental heart must be separated
from the others by a valve. To make the matter wholly clear, we may
observe an insect heart pulsating, and this is best seen in one of
its middle chambers. This chamber expands (simply by the relaxation
of its circular muscles), the ostia, also, consequently open, and a
given quantity of blood is drawn in from the pericardial cavity.
What now would happen after the succeeding contraction if there were
no valves between? The blood would not flow forwards, but seek a way
out backwards.

“But, in fact, the valve of the hinder chamber, at this time, closes
itself, while, by the simultaneous expansion of the anterior ones,
their door opens, and this section of the heart, at the same time,
causes a sucking in of the contents of the posterior chamber. This
phenomenon is repeated, in the same way, from chamber to chamber,
which also acts alternately as ventricle and auricle, or by a
sucking and pumping action. One is involuntarily reminded of the
ingenious manipulation by which, by the alternate opening and
shutting of the flood-gates, a vessel is carried along a canal.

“This wave-like motion of an insect’s heart also has the advantage
that, just before a pulse-wave has reached the chambers farthest in
front, the hinder ones are already prepared for the production of a
second, for, as a matter of fact, often 60, and even 100, and, in
very agile insects, 150, waves pass, in a single minute, through the
series of chambers, which make it very difficult to follow the
flowing of their waves.” (Graber.)

=The propulsatory apparatus.=—But the heart itself is only a part of
the entire propulsatorial apparatus to which belongs the following
contrivance, the nature of which has been worked out by Graber.

Under the dorsal vessel is stretched a sort of roof-like diaphragm,
_i.e._ a membrane, arched like the dorsal wall of the hind-body
which is attached, in a peculiar way, to the sides of the body. The
best idea can be gained by a cross-section through the entire body
(Fig. 374): _H_ is the true dorsal vessel; _S_, the diaphragm. A
surface view is seen at 373, _C_, _b_, where it appears as a plate
with the edge regularly curved outwards on each side. Its precise
mode of working is thus: from each dorsal band of the sides of the
abdomen arises a pair of muscles spreading out fan-like, and
extending to the heart, so that the fibres of one side pass directly
over to those of the other, often splitting apart, or, between the
two, extends outwards a perforated, thin web, like an elastic,
fibrous sheet (Fig. 373, _A_, _a_), with numerous perforations,
forming a diaphragm.

[Illustration:

FIG. 374.—Diagram of transverse section of pericardial sinus of
_Ædipoda cœrulescens_: _H_, heart; _s_, septum; _m_, muscles,—the
upper suspensory, the lower alary.—After Graber, from Sharp. (See
also Fig. 377.)
]

Graber has thus explained the action of the pericardial diaphragm
and chamber, as freely translated by Miall and Denny: “When the
alary muscles contract, they depress the diaphragm, which is arched
upwards when at rest. A rush of blood towards the heart is thereby
set up, and the blood streams through the perforated diaphragm into
the pericardial chamber. Here it bathes a spongy or cavernous tissue
(the fat-cells), which is largely supplied with air-tubes, and
having been thus aerated, passes immediately forwards to the heart,
entering it at the moment of diastole, which is simultaneous with
the sinking of the diaphragm.”

In the cockroach, however, Miall and Denny think that the facts of
structure do not altogether justify this explanation: “The fenestræ
of the diaphragm are mere openings without valves. The descent of a
perforated non-valvular plate can bring no pressure to bear upon the
blood, for it is not contended that the alary muscles are powerful
enough to change the figure of the abdominal rings.... The diaphragm
appears to give mechanical support to the heart, resisting pressure
from a distended alimentary canal, while the sheets of fat-cells, in
addition to their proper physiological office, may equalize small
local pressures, and prevent displacement. The movement of the blood
towards the heart must (we think) depend, not upon the alary
muscles, but upon the far more powerful muscles of the abdominal
wall, and upon the pumping action of the heart itself.”

“The peculiar office,” says Graber, “performed by the heart has
already been stated. It is nothing more than a regulator; than an
organ for directing the blood in a determinate course in order that
this may not wholly stagnate, or only be the plaything of a force
acting in another way, as, for example, through that afforded by the
body-cavity and the inner digestive canal. At regular intervals a
portion of the blood is sucked through the same, and then by means
of the anterior supply tube it is pushed onward into the head,
whence it passes into the cavities of the tissues. The different
conditions of tension under which the mass of blood stands in the
different regions of the body then causes a farther circulation.
Besides this, the blood passes through separate smaller pumping
apparatuses, and through vessel-like modifications of cavities, also
through hollow spaces between the muscles, as, for example, in the
appendages where a regular backward and forward flow of the blood,
especially in the limbs, wings, antennæ, and certain abdominal
appendages takes place. Here and there may occasionally occur a
narrow place where the flow of blood is obstructed by the
accumulation of the blood corpuscles, causing a considerable
stagnation.” (Graber.)

[Illustration:

FIG. 375.—_Libellula depressa_, opened from the back, showing the
nervous cord (_b_{1}_-_b_{3}_, thoracic, _h_{1}_-_h_{7}_, abdominal,
ganglia), also the furrow-like ventral sinus closed by a muscular
diaphragm.
]

[Illustration:

FIG. 376.—_A_, part of the ventral furrow of _Libellula depressa_ more
highly magnified: _a_, a sternal plate (urite); _c_, the septum
stretched over it, at _s_ in a relaxed or collapsed state; _b_ and
_d_, the wing-like, sternal processes from which the muscular
bundles of the diaphragm arise. _B_, same in Acridium.
]

[Illustration:

FIG. 377.—Diagrammatic section of the abdomen of _Acridium
tartaricum_, showing the ventral septum (_i_, _p_, _l_) contracted,
and (_i_, _k_, _l_) stretched out; _oh_, rib-like lateral processes
of the urite; _f_, ganglia; _b_, heart, with its suspensorium (_a_);
_c_, fat tissue in the pericardial tissue sinus; _d_, dorsal septum
or diaphragm contracted. _q_, extended; _g_, fat-body; _e_, muscular
part of diaphragm; _no_, expiration, _hm_, inspiration, muscle.—This
and Figs. 375, 376, after Graber.
]

=The supraspinal vessel.=—In many insects there is a ventral heart
acting on the heart’s blood as an aspirator, or more correctly a ventral
sinus lying on the nervous cord, and closed by a pulsating diaphragm.
This was discovered by Réaumur in the larva of a fly, and by Graber in
the dragon-fly and locusts (Acrydiidæ). A glance at Figs. 375 and 376
will save a long description. The ventral wall forms a furrow, and
between its borders (Fig. 377, _e_) extends the diaphragm. During the
contraction of the muscles—and this, here, acts from before
backwards—the membrane rises up and makes a cavity for the blood, which
passes backwards over the nervous cord. The dorsal and ventral sinuses
together thus bring about a closed circulation.

It thus appears that the insects are well provided with the means of
distribution of their nutritive fluid, and that the blood is kept
continually fresh and rich in oxygen. (Graber.)

=The aorta.=—While the heart is mostly situated within the abdomen, it
is continued into the thorax and the head as a simple, non-pulsating
tube, called the aorta. In Sphinx the aorta, as described by Newport,
begins at the anterior part of the 1st abdominal segment, where it bends
downwards to pass under the metaphragma and enter the thorax; it then
ascends again between the great longitudinal dorsal muscles of the
wings, and passes onwards until it arrives at the posterior margin of
the pronotum; it then again descends and continues its course along the
upper surface of the œsophagus, with which it passes beneath the brain,
in front of which and immediately above the pharynx, it divides into two
branches, each of which subdivides. Newport, however, overlooked a
thoracic enlargement of the aorta called by Burgess the “aortal chamber”
(Fig. 310, _a_, _c_).

[Illustration:

FIG. 378.—_A_, last three abdominal segments and bases of the three
caudal processes of _Cloëon dipterum_: _r_, dorsal vessel; _kl_,
ostia; _k_, special terminal chamber of the dorsal vessel with its
entrance _a_; _b_, blood-vessel of the left caudal process. _B_,
26th joint of the left caudal appendage from below: _b_, a portion
of the blood-vessel; _o_, orifice in the latter.—After Zimmermann,
from Sharp.
]

“In Sphinx and _Vanessa urticæ_, immediately after the aorta has
passed beneath the cerebrum, it gives off laterally two large
trunks, which are each equal in capacity to about one-third of the
main vessel. These pass one on each side of the head, and are
divided into three branches which are directed backwards, but have
not been traced farther in consequence of their extreme delicacy.
Anterior to these trunks are two smaller ones which appear to be
given to the parts of the mouth and antennæ, and nearer the median
line are two others which are the continuations of the aorta. These
pass upwards, and are lost in the integument. The whole of these
parts are so exceedingly delicate that we have not, as yet, been
able to follow them beyond their origin at the termination of the
aorta, but believe them to be continuous, with very delicate,
circulatory passages along the course of the tracheal vessels. It is
in the head alone that the aorta is divided into branches, since,
throughout its whole course from the abdomen, it is one continuous
vessel, neither giving off branches, nor possessing lateral muscles,
auricular orifices, or separate chambers.” (Newport, art. Insecta,
p. 978.)

Dogiel observed in the transparent larva of _Corethra plumicornis_ that
the aorta extends only to the hinder border of the brain. Here it
divides into two lamellæ, each of which independently extends farther
on. One lamella is seen under the brain and under the eye, the other
reaches near the eye. The lamellæ are tied to the integument by threads.
At the point of division of the aorta is an opening. (Kolbe.)

True blood-vessels appear to exist in the caudal appendages of the
May-flies, as the heart appears to divide and pass directly into
them (Fig. 378). The last chamber of the heart diminishes in size at
the end of the body, and then divides into three delicate tubular
vessels which pass into the three caudal appendages, and extend to
the end of each one, along the upper side. While the valves of the
heart, in all insects, are directed anteriorly because the blood
flows from behind, in the larva of the Ephemeridæ the valves of the
last chamber of the heart are directed backwards, because from this
chamber the blood flows in the opposite direction, _i.e._ into the
caudal appendages. During the contraction of the heart, the
elongated section of the same in the last abdominal segment receives
a part of the mass of blood contained in the last chamber, which is
driven by independent contractions into the caudal appendages. These
vessels have openings before the end through which the blood enters
into the cavity of the appendages, and can also pass back, in order
to be taken up by the body cavity. It is possible that these
blood-vessels stand in direct relation to respiration. (Zimmermann,
Creutzburg, in Kolbe, p. 544.)

=The pericardial cells.=—Along the heart, on both sides, occur the
so-called pericardial cells, which differ from the fat-cells, and
also the peritracheal cells of Frenzel, and are mostly arranged in
linear series, which have a close relation to the circulation of the
blood. In the larva of Chironomus, they lie in groups; in that of
Culex, they are arranged segmentally. In caterpillars, these
pericardial cells are not situated in the region of the heart, but
are arranged linearly on the side, and form a network of granulated
cells situated between the fat-bodies. Other rows of these cells are
situated near the stigmata and the main lateral tracheæ. (Kolbe.)

According to Kowalevsky, the pericardial cells, and the
garland-shaped, cellular cord consist of cells, whose function it is
to purify the blood, and to remove the foreign or injurious matters
mingled with the blood.

[Illustration:

FIG. 379.—Diagram of the circulatory organs in the head of the
cockroach, seen from above: _A_, ampulla; _V_, antennal vessel;
_M_, chief muscular cord; _m_, muscular band; _Bs_, wall of the
blood sinus; _am_, opening of the aorta (a); _rg_, anterior
sympathetic or visceral ganglion; _hg_, hinder visceral ganglion;
_F_, _F_, facetted eyes; _o_, vestigial ocellus; _G_, _G_, brain;
_S_, œsophagus.—After Pawlowa.
]

=Ampulla-like blood circulation in the head.=—In the head of the
cockroach occurs, according to Pawlowa, a contractile vascular sac
at the base of each antenna. The cavity has a valvular communication
with the blood space below and in front of the brain, and
muscle-fibres effect systole and diastole. Each sac is beyond doubt
an independently active part of the circulatory system. These organs
also occur in Locusta and other Acrydiidæ, and Selvatico has
described similar structures in _Bombyx mori_ and certain other
Lepidoptera.

=Pulsatile organs of the legs.=—Accessory to the circulation is a
special system of pulsatile organs in the three pairs of legs of
Nepidæ, generally situated in the tibia just below its articulation
with the femur, but in the fore legs of Ranatra, in the clasp-joint
or tarsus, just below its articulation with the tibia. First
observed by Behn (1835), Locy has studied the organ (Fig. 380) in
Corixa, Notonecta, Gerris, besides the Nepidæ. It is a whip-like
structure attached at both ends, with fibres extending upward and
backward to the integument of the leg, separate from the muscular
fibres and does not involve them in its motions, and is not affected
by the muscles themselves. “As the blood-corpuscles flow near the
pulsating body they move faster, and around the organ itself there
is a whirlpool of motion.” The beating of these organs aids the
circulation in both directions, and when the motion ceases, the
blood-currents in the legs stop; the rate of the pulsating organ is
always faster than that of the heart, and the action is automatic.

[Illustration:

FIG. 380.—Pulsating organs in Hemiptera: _A_, Belostoma nymph, _B_,
legs of Corixa. _C_, Ranatra, adult, to show the exceptional
position of the pulsating organ in the fore legs. _D_, pulsatile
organ in tibia of Ranatra.—After Locy.
]

_b._ The blood

The blood of insects, as in other invertebrates, differs from that of
the higher animals in having no red corpuscles. It is a thin fluid, a
mixture of blood (serum) and chyle, usually colorless, but sometimes
yellowish or reddish, which contains pale amœboid corpuscles
corresponding to the white corpuscles (leucocytes) of the vertebrates,
though they are relatively less numerous in the blood of insects. The
yellow fluid expelled from the joints of certain beetles (Coccinella,
Timarcha, and the Meloidæ) is, according to Leydig, only the serum of
the blood. In phytophagous insects the blood is colored greenish by the
chlorophyll set free during digestion. The blood of _Deilephila
euphorbia_ is colored an intense olive-green, and that of _Cossus
ligniperda_ is pale yellow. (Urech.) The blood of case-worms
(Trichoptera) is greenish. In some insects it is brownish or violet. The
serum is the principal bearer of the coloring material, yet Graber has
shown that in certain insects the corpuscles are more or less beset with
bright yellow or red fat-globules, so as to give the same hue to the
blood.

=The leucocytes.=—The corpuscles are usually elongated, oval, or
flattened oat-shaped, with a rounded nucleus, or are often amœbiform;
and they are occasionally seen undergoing self-division. When about to
die the corpuscles become amœbiform or star-shaped. (Cattaneo.) Their
number varies with the developmental stage of the insect, and in larvæ
increases as they grow, becoming most abundant shortly before pupation.
The blood diminishes in quantity in the pupal stage, and becomes still
less abundant in the imago. (Landois.) The quantity also varies with the
nutrition of the insect, and after a few days’ starvation nearly all the
blood is absorbed. Crystals may be obtained by evaporating a drop of the
blood without pressure; they form radiating clusters of pointed needles.
The freshly drawn blood is slightly alkaline. (Miall and Denny.)

The size of the corpuscles has been ascertained by Graber, who found
that the diameter of the circular blood-disks of the leaf-beetle,
_Lina populi_, is 0.006 mm.; of _Cetonia aurata_ and _Zabrus
gibbus_, 0.008 to 0.01 mm.; and those of certain Orthoptera
_(Decticus verrucivorus_, _Ephippiger vitium_ and _Œdipoda
cœrulescens_), 0.011 to 0.014 mm. The longest diameter of the
elongated corpuscles of _Carabus cancellatus_ is 0.008 mm.; of
_Gryllus campestris_, _Locusta viridissima_, _Cossus ligniperda_,
_Sphinx ligustri_ (pupa), and others, 0.008 to 0.01 mm.; of
_Caloptenus italicus_, _Saturnia pyri_, _Anax formosus_, and others,
0.011 to 0.014 mm.; of _Ephippiger vitium_, _Œdipoda cœrulescens_,
_Pezotettix mendax_, _Zabrus gibbus_, _Phryganea_, and others, 0.012
to 0.022 mm.; in _Stenobothrus donatus_ and _variabilis_, 0.012 to
0.035 mm. The largest known are those of _Melolontha vulgaris_,
which measure from 0.027 to 0.03 mm.

[Illustration:

FIG. 381.—Blood corpuscles, or leucocytes, of insects: _A_, _a-g_, of
_Stenobothrus dorsatus_ (the same forms occur in most Orthoptera and
in other insects). _B_, _a_, leucocyte of the same insect with the
nucleus brought out by ether; _b_, another of serpentine shape. _C_,
leucocytes of the same insect after a longer stay in ether. _D_,
leucocytes of the same after being in glycerine 14 days.—After
Graber.
]

As regards the nature of the corpuscles, Graber concludes that they are
more like the cells of the fat-bodies than genuine cells. That they are
not true cells is shown by the fact that after remaining in their normal
condition a long time they finally coalesce and form cords. After
shrivelling, or after the blood has been subjected to different kinds of
treatment, the nucleus is clearly brought out (Fig. 381).

Besides the blood corpuscles there have been detected in the blood round
bodies which are regarded as fat-cells. They are circular, and for the
most part larger than the blood corpuscles, have a sharp, even, dark
outline, and an invariably circular nucleus. (Kolbe.)

The blood of Meloe, besides the amœboid corpuscles, according to
Cuénot, contains abundant fibrinogen, which forms a clot; a pigment
(uranidine), which is oxidized and precipitated when exposed to the
air; a dissolved albuminoid (hæmoxanthine), which has both a
respiratory and nutritive function; and, finally, dissolved
cantharidine.

The corpuscles arise from tissues which are very similar to the
fat-bodies, and which, at given times, separate into cells. The
position of these tissues is not always the same in different
insects. In caterpillars, they occur in the thorax, near the germs
of the wings; in the saw-flies (Lyda), in all parts of the thorax
and abdomen; in larval flies (Musca), in the end of the abdomen,
just in front of the large terminal stigmata. The place where the
blood corpuscles are formed is usually near, or in relation with,
the fat-bodies. But while the fat-bodies mostly serve as the
material for the formation of the blood-building tissues, in
caterpillars the tracheal matrix also, and, in dipterous larvæ, the
hypodermis serve this purpose. (Cæsar Schaeffer in Kolbe. See also
Wielowiejski, Ueber das Blutgewebe.)

Other substances occur in the blood of insects. Landois (1864)
demonstrated the existence of egg albumen, globulin, fibrin, and
iron in the blood of caterpillars. Poulton found that the blood of
caterpillars often contained chlorophyll and xanthophyll derived
from their food plants. A. G. Mayer has recently found that the
blood (hæmolymph) of the pupæ of Saturniidæ (_Callosamia promethea_)
contains egg albumin, globulin, fibrin, xanthophyll, and
orthophosphoric acid, and Oenslager has determined that iron,
potassium, and sodium are also present. (Mayer.)

_c._ The circulation of the blood

Every part of the body and its appendages is bathed by the blood, which
circulates in the wings of insects freshly emerged from the nymph or
pupal state, and even courses through the scales of Lepidoptera, as
discovered by Jaeger (Isis, 1837).

In describing the mechanism of the heart we have already considered in a
general way the mode of circulation of the blood.

The heart pumping the blood into the aorta, the nutritive fluid passes
out and returns along each side of the body; distinct, smaller streams
passing into the antennæ, the legs, wings (of certain insects), and into
the abdominal appendages when they are present. All this may readily be
observed in transparent aquatic insects, such as larval Ephemeræ,
dragon-flies, etc., kept alive for the purpose under the microscope in
the animalcule box.

Carus, in 1827, first discovered the fact of a complete circulation of
the blood, in the larva of Ephemera. He saw the blood issuing in several
streams from the end of the aorta in the head and returning in currents
which entered the base of the antennæ and limbs in which it formed
loops, and then flowing into the abdomen, entered the posterior end of
the heart. Wagner (Isis, 1832) confirmed these observations, adding one
of his own, that the blood flows backward in two venous currents, one at
the sides of the body and intestine, and the other alongside of the
heart itself, and that the blood not only entered at the end of the
heart, but also at the sides of each segment, at the position of the
valves discovered by Straus-Dürckheim.

Newport maintains that the course of the blood is in any part of the
body, as well as in the wings, almost invariably in immediate connection
with the course of the tracheæ, for the reason that “the currents of
blood in the body of an insect are often in the vicinity of the great
tracheal vessels, both in their longitudinal and transverse direction
across the segments.”

The circulation of the blood in the wings directly after the exuviation
of the nymph or pupa skin, and before they become dry, has been proved
by several observers. As stated by Newport, the so-called “veins” or
“nervures” of the wings consist of tracheæ lying in a hollow cavity, the
peritracheal space being situated chiefly under and on each side of the
trachea.

[Illustration:

FIG. 382.—Circulation of the blood in hind wing of _Periplaneta
orientalis_: the arrows indicate the usual direction of the blood
currents.—After Moseley.
]

Newport gives the following summary of the observations of the early
observers, to which we add the observations of Moseley. “A motion of
the fluids has been seen by Carus in wings of recently developed
Libellulidæ, _Ephemera lutea_ and _E. marginata_, and _Chrysopa
perla_; among the Coleoptera, in the elytra and wings of _Lampyris
italica_ and _L. splendidula_, _Melolontha solstitialis_ and
Dytiscus.” Ehrenberg saw it in Mantis, and Wagner in the young of
_Nepa cinerea_ and _Cimex lectularius_. Carus detected a circulation
in the pupal wings of some Lepidoptera, and Bowerbank witnessed it
in a Noctuid (_Phlogophora meticulosa_); Burmeister observed it in
_Eristalis tenax_ and _E. nemorum_, and Mr. Tyrrel in _Musca
domestica_, but it has not been observed in the wings of
Hymenoptera.

Bowerbank observed that in the lower wing of _Chrysopa perla_ the
blood passes from the base of the wing along the costal,
post-costal, and externo-medial veins, outwards to the apex of the
wing, giving off smaller currents in its course, and that it returns
along the anal vein to the thorax. He found that the larger veins,
1⁄408 in. in diameter, contained tracheæ which only measured 1⁄2222
of an inch in diameter; but in others the tracheæ measured 1⁄1340,
while the cavity measured only 1⁄500 of an inch. He states, also,
that the tracheæ very rarely give off branches while passing along
the main veins, and that they lie along the canals in a tortuous
course. (Newport, art. Insecta, p. 980.)

Bowerbank, also, in his observations on the circulation in the wings
of Chrysopa, “used every endeavor to discover, if possible, whether
the blood has proper vessels, or only occupied the internal cavities
of the canals; and that he is convinced that the latter is the case,
as he could frequently perceive the particles not only surrounding
all parts of the tracheæ, and occupying the whole of the internal
diameter of the canals, but that it frequently happens that gl